/ 


THE  UNIVERSITY 

OF  ILLINOIS 

LIBRARY 

NATURAL  HISTORY  SURVEY 


ViTim 


a 


books.  ^   overdue 

U.  of^.  Library 


ILLINOIS   BIOLOGICAL 
MONOGRAPHS 

Vol.  V  July-October,  1919  Nos.  3  and  4 


Editorial  Committee 


Stephen  Alfred  Forbes  William  Trelease 

Henry  Baldwin  Ward 


PUBUSHEO  UNDER  THE 

Auspices  or  the  Graduate  School  by 
THE  University  or  Illinois 


Copyright,  1920  by  the  UNivERsrry  o»  Ilunots 
Distributed  December  31, 1920. 


STUDIES  ON  MYXOSPORIDIA 

A  SYNOPSIS  OF  GENERA  AND  SPECIES 
OF  MYXOSPORIDIA 


WITH  25  PLATES  AND  2  TEXTFIGURES 


BY 

ROKUSABURO  KUDO 


Contributions  from  the 

Zoological  Laboratory  of  the  University  of  Illinois 

No.  158 


TABLE  OF  CONTENTS 

Introduction 7 

General  Remarks  on  Recent  Observations 8 

Myxosporidia  Recorded  in  the  Present  Paper  (List  I) 9 

Distribution  of  Myxosporidia 13 

A.  Geographical  Distribution 13 

B.  Distribution  of  Myxosporidia  in  Animals 25 

C.  Distribution  of  Myxosporidia  in  the  Organs  of  the  Host 37 

D.  The  Efifect  of  Environment  on  the  Organal  Distribution  of  Mj^osporidia  in 

Hosts 44 

The  Spore 47 

Definition  of  Terms  Used  for  Descriptions 50 

Classification  of  Myxosporidia 52 

Descriptions  of  Genera  and  Species 60 

Suborder  Eurysporea '.  60 

Family  Ceratomyxidae 60 

Genus  Leptotheca 60 

Genus  Ceratomyxa 65 

Genus  Myxoproteus 81 

Genus  Wardia 82 

Genus  Mitraspora 84 

Suborder  Sphaerosporea 86 

Family  Chloromyxidae 87 

Genus  Chloromyxum 87 

Family  Sphaerosporidae 99 

Genus  Sphaerospora 100 

Genus  Sinuolinea 104 

Suborder  Platysporea 106 

Family  Myxidiidae 106 

Genus  Myxidium 107 

Genus  Sphaeromyxa 118 

Genus  Zschokkella 122 

Family  Myxosomatidae 123 

Genus  Myxosoma 123 

Genus  Lentospora 125 

Family  Myicobolidae 128 

Genus  Myxobolus ^>!-^^. 128 

Genus  Henneguya 158 

Genus  Hoferellus 173 

Myxosporidia  Genera  et  Species  Incertae 174 


6                                   ILUNOJS  BIOLOGICAL  MONOGRAPHS  [244 

K^s  to  the  Genera  and  Species  of  Myxosporidia 178 

Key  to  the  Genera  of  Myxosporidia 178 

Key  to  the  Species 179 

Summary 196 

Appendix:  New  Myxosporidia  from  Australia 197 

Bibliography 200 

General  Explanation  of  Figures 210 

Explanation  of  Plates 212 

Index 261 


245j  STUDIES  ON  MYXOSPORIDIA—KUDO 


INTRODUCTION 

Ten  years  have  elapsed  since  Auerbach  (1910)  published  Die  Cnido- 
sporidien  in  which  he  gave  a  synopsis  of  the  genera  and  species  of  Myxos- 
poridia  known  up  to  that  time.  During  this  period  new  genera  and  a 
number  of  new  species  have  been  added  to  the  list  of  this  particular  group 
of  parasitic  protozoa  from  the  various  parts  of  the  world.  It  is,  therefore, 
desirable  to  have  a  complete  monographic  work  including  all  the  forms 
reported  up  to  the  present  time. 

The  main  objects  of  the  present  paper  are:  1)  to  describe  a  new  genus 
and  a  number  of  new  species  which  have  come  under  the  observation  of  the 
writer;  2)  to  collect  all  the  genera  and  species  recorded  by  various  authors; 
3)  to  propose  a  new  classification  by  which  some  of  the  confusion  now 
existing  may,  probably,  be  avoided;  4)  to  show  the  geographical,  zoological 
and  organal  distribution  in  the  light  of  more  recent  observations;  and 
5)  to  present  a  complete  list  of  the  names  of  the  hosts  in  which  Myxospori- 
dia  occur. 

The  writer  believes  himself  to  be  in  possession  of  as  complete  references 
as  possible  under  present  conditions.  However,  he  may  be  unaware  of 
some  works  which  have  not  reached  him  owing  to  the  war. 

The  Myxosporidia  recorded  by  Labbe  (1899)  are  arranged  in  almost  the 
same  order  as  that  author  listed  them,  with  some  slight  change  such  as 
placing  the  type  species  at  the  front  of  each  genus  or  removing  a  few  species 
to  other  genera,  while  those  species  which  have  been  described  since  1898 
are  arranged  chronologically,  no  matter  whether  names  are  given  the 
species  or  not. 

Some  of  the  references  are  omitted,  especially  when  they  can  be  found 
in  Gurley  (1894),  Thelohan  (1895),  Labbe  (1899),  or  Auerbach  (1910). 
The  description  of  each  species  is  given  according  to  the  first  observer. 
The  observations  of  subsequent  investigators  are  then  mentioned  in  the 
second  place. 

Each  species  is  described  according  to  the  following  scheme: 

1)  Specific  name 

2)  Synonyms  and  literature 

3)  Habitat,  including  the  locaHty  and  the  date  of  observation 

4)  Vegetative  form  \^ 

5)  Spore  ^^"^ 

6)  Remarks 

I  wish  to  express  my  appreciation  to  Professor  Henry  B.  Ward  whose 
kindness  has  made  the  completion  of  this  paper  possible. 


ILUNOJS  BIOLOGICAL  MONOGRAPHS  (246 


GENERAL  REMARKS  ON  RECENT  OBSERVATIONS 

The  total  number  of  species  of  Myxosporidia  reported  up  to  date  and 
described  in  the  following  pages,  excluding  12  ambiguous  forms,  reaches 
237  of  which  125  are  species  which  have  been  observed  since  1910,* 
The  distribution  of  these  new  forms  is  as  follows: 

Africa.. 6  species 

Asia 23  species 

Australia 1  species 

Europe 31  species 

North  America 63  species 

South  America 1  species 

Thus,  the  majority  of  the  species  were  observed  in  other  lands  than 
Europe,  nearly  half  being  recorded  from  North  American  waters.  It  is  not 
hard  to  anticipate  from  the  observations  made  by  Awerinzew,  Davis, 
Kudo,  Mavor,  Johnston  and  Bancroft,  and  others,  that  further  investiga- 
tions on  the  parasites  in  the  localities  where  the  study  of  the  protozoa 
under  consideration  was  neglected,  will  bring  out  not  only  new  and  inter- 
esting forms  which  will  be  quite  different  from  the  comparatively  well 
studied  European  species,  but  also  many  important  facts  that  will  clear 
unknown  or  doubtful  phases  concerning  the  life  history  and  structure  of 
Myxosporidia. 

*  Three  species  are  included  here  which  have  been  described  (in  Nipponese)  by  Miyairi' 
in  1909. 


247] 


STUDIES  ON  MYXOSPORJDIA—KUDO 


MYXOSPORIDIA  RECORDED  IN  THE  PRESENT  PAPER 

LIST  I 

Order    MYXOSPORIDIA    Butschli 

I  Suborder    EURYSPOREA    nom.  nov.     (see  page  56) 

I  Family    CERATOMYXIDAE    Doflein 


Genus  1    LEPTOTHECA    Th61ohan 
[15  species] 

1)  L.  agilis  Th^lohan  (type  species) 

2)  L.  elongata  Th61ohan 

3)  L.  polymorpha  (Th61.)  Labbe 

4)  L.  parva  Th61ohan 

5)  L.  renicola  Th61ohan 

6)  L.  hepseti  Th61ohan 

7)  L.  perlata  (Gurley)  Labb6 

8)  L.  sp.  Awerinz«w 

9)  L.  macrospora  Auerbach 

10)  L.  informis  Auerbach 

11)  i.  longipes  Auerbach 

12)  L.  fusiformis  Davis 

13)  L.  scissura  Davis 

14)  L.  lobosa  Davis 

15)  L.  glomerosa  Davis 

Genus  2     CERATOMYXA    Thelohan 
[35  species] 

1)  C.  arcuata  Th6Iohan  (type  species) 

2)  C.  sphaertdosa  Thelohan 

3)  C.  pallida  Th61ohan 

4)  C.  globurifera  Thelohan 

5)  C.  appendicidata  Thelohan 

6)  C.  truncata  Thelohan 

7)  C.  retictdaris  Thelohan 

8)  C.  inaequalis  Doflein 

9)  C.  linospora  Doflein 

10)  C.  ramosa  Awerinzew 

11)  C.  drepanopsettae  Awerinzew 

12)  C.  tylosuri  Awerinzew 

13)  C.  (?)  spari  Awerinzew 

14)  C.  sp.  (?)  Awerinzew 

15)  C.  sp.  (?)  Awerinzew 

16)  C.  acadiensis  Mavor 


17) 

C.  sp.  Georgevitch 

18) 

C.  coris  Georgevitch 

19) 

C.  herouardi  Georgevitch 

20) 

C.  mesospora  Davis 

21) 

C.  sphairophora  Davis 

22) 

C.  taenia  Davis 

23) 

C.  atlenuata  Davis 

24) 

C.  recurvata  Davis 

25) 

C.  lunata  Davis 

26) 

C.  abbreviata  Davis 

27) 

C.  flagellifera  Davis 

28) 

C.  agglomerata  Davis 

29) 

C.  anwrpha  Davis 

30) 

C.  nwnospora  Davis 

31) 

C.  streptospora  Davis 

32) 

C.  aggregata  Davis 

33) 

C.  undtdata  Davis 

34) 

C.  navicularia  Davis 

35) 

C.  spinosa  Davis 

Genus  3    MYXOPROTEUS    Doflein 

[3  species] 

1) 

M.  ambiguus  (Thelohan)  Doflein  (ty 

species) 

2) 

M.  cordifortnis  Davis 

3) 

M.  cornutus  Davis 

Genus  4    WARDIA    nov.  gen. 
[2  species] 

1)  W.  ovinocua  nov.  spec,  (type  species) 

2)  W.  ohlmacheri  (Gurley)  Kudo 

Genus    5  MITRASPORA    Fujita    emend. 
Kudo    [3  species] 

1)  M.  cyprini  Fujita  (type  species) 

2)  M.  caudata  (Parisi)  Kudo 

3)  M.  elongata  nov.  spec. 


10 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[248 


II  Suborder    SPHAEROSPOREA    nom.  nov.  (see  page  57) 
I  FamUy    CHLOROMYXIDAE    Thfilohan 


Genus  1     CHLOROMYXUM     Mingazzini 


1)  C. 

2)  C. 

3)  C. 

4)  C. 

5)  C. 

6)  C. 

7)  C. 

8)  C. 

9)  C. 
10)  C. 


[22  species] 
leydigi  Mingazzini  (type  species) 
caudatum  Tli61ohan 
quadratutn  Th6Iohan 
fluviatUe  Th61ohan 
mucronatutn  Gurley 
diploxys  (Gurley)  Th61ohan 
protei  Joseph 
truttae  lAgtr 
cristatum  L^ger 
dubiutn  Auerbach 


11) 
12) 
13) 
14) 
15) 
16) 

17)  C. 

18)  C. 

19)  C. 

20)  C. 

21)  C. 

22)  C. 


sp.  Awerinzew 
thymaUi  Lebzelter 
koi  Fujita 

magnum  Awerinzew 
fundtdi  Hahn 
misgurni  Kudo 
fujitai  Kudo 
clupeidae  Hahn 
granulosum  Davis 
trijugum  nov.  spec. 
catostomi  nov.  sp>ec. 
joarJx  nov.  spec. 


II  Fanuly    SPHAEROSPORIDAE    Davis 


Genus  1    SPHAEROSPORA  Th^lohan 
[10  species] 

1)  S.  divergens  Th61ohan  (type  species) 

2)  S.  elegans  Thfilohan 

3)  S.  rostrata  Th^lohan 

4)  5.  masovica  Cohn 

5)  S.  platessae  Woodcock 

6)  S.  angulata  Fujita 

7)  S.  sp.  Davis 

8)  5.  polymorpha  Davis 


9)  S.  (?)  sp.  Southwell  et  Prashad 
10)  S.  carassii  nov.  spec. 

Genus  2    SINUOLINEA    Davis 
[5  species] 

1)  5.  dimorpha  Davis  (type  species) 

2)  S.  capsularis  Davis 

3)  S.  arborescens  Davis 

4)  S.  opacita  Davis 

5)  S.  brachiophora  Davis 


ni  Suborder    PLATYSPOREA    nom.  nov.     (see  page  57) 
I  FamUy    MYXIDIIDAE    Th6lohan 


Genus  1    MYXIDIUM    BUtschli 
[26  species] 

1)  M.  lieberkiihni  Btitschli  (type  species) 

2)  M,  incurvatum  Th61ohan 

3)  M.  sphaericum  Th61ohan 

4)  M.  histophUum  Th^lohan 

5)  M.  ^.  Gurley 

6)  M.  danilewskyi  Laveran 

7)  M.  giganteum  Doflein 

8)  M.  barbatulae  C6pfede 

9)  M.  giardi  Cepede 

10)  M.  pfdferi  Auerbach 

11)  M.  inflatum  Auerbach 

12)  M.  bergense  Auerbach 

13)  M.  procerum  Auerbach 

14)  M.  mackiei  Bosanquet 

15)  M.  macrocapstdare  Auerbach 

16)  M.  sp.  Awerinzew 

17)  M.  depressum  Parisi 

18)  M.  oviforme  Parisi 

19)  M.  anguillae  Ishii 

20)  M.  sp.  Mavor 


21)  M.  gadi  Georg6vitch 

22)  M.  glutinosum  Davis 

23)  M.  phyllium  Davis 

24)  M.  striatum  Cunha  et  Fonseca 

25)  M.  kagayamai  nov.  spec. 

26)  M.  americanum  nov,  spec. 

Genus  2    SPHAEROMYXA    Thaohan 

[7  species] 

1)  5.  balbianii  Th61ohan  (type  species) 

2)  S.  immersa  (Lutz)  Th61ohan 

3)  S.  incurvata  Doflein 

4)  S.  sabrazesi  Laveran  et  Mesnil 

5)  S.  hellandi  Auerbach 

6)  S.  exneri  Awerinzew 

7)  S.  gasterostei  Georgevitch 

Genus  3    ZSCHOKKELLA    Auerbach 
[4  spedes] 

1)  Z.  hildae  Auerbach  (type  species) 

2)  Z.  nova  Klokacewa 

3)  Z.  acheilognathi  Kudo 

4)  Z.  globulosa  Davis 


249] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


If 


n  Fanuly    MYXOSOMATIDAE    Poche 


Genus  1    MYXOSOMA    Thdlohan 
[3  species] 

1)  M.  dujardini  Th61ohan  (type  species) 

2)  M.  (?)  lobatum  Nemeczek 

3)  M./unduli  Kudo 


1 

2 

3 

4; 

5 

6 

7 

8 

9; 

10 

11 

12 

13 

14; 

15 

16 

17 

18 

19 

20: 

21 

22 

23 

24 

2^: 

26; 
27 
28 
29 
30; 
31 
32 
33 
34; 
35 
36 
37 
38 


III  Family    MYXOBOL 


Genus  1    MYXOBOLUS    BUtschli 
[63  species] 
M.  mulleri  Biitschli  (type  species) 
M.  piriformis  Thfilohan 
M.  unicapstdatus  Gurley 
M.  fuhrmanni  Auerbach 
M.  octdi-leucisci  Trojan 
M.  toyamai  Kudo 
M.  notattis  Mavor 
M.  sp.  Kudo 

M.  rohitae  Southwell  et  Prashad 
M.  seni  Southwell  et  Prashad 
M.  tnisgurni  nov.  spec. 
M.  pfeiferi  Th61ohan 
M.  inaequalis  Gurley 
M.  dispar  Th61ohan 
M.  ellipsoides  Th^Iohan 
M.  exiguus  Thelohan 
M.  oviformis  Th61ohan 
M.  lintoni  Gurley 
M.  globosus  Gurley 
M.  ohlongus  Gurley 
M.  transovalis  Gurley 
M.  obesus  Gurley 
M.  cycloides  Gurley 
M.  sphaeralis  Gurley 
M.  anurus  Cohn 
M.  sp.  Gurley 
M.  sp.  Gurley 
M.  sp.  Gurley 
M.  cyprini  Doflein 
M.  neurohius  Schuberg  et  Schroder 
M.  aeglefini  Auerbach 
M.  gigas  Auerbach 
M.  volgensis  Reuss 
M.  scardinii  Reuss 
M.  physophilus  Reuss 
M.  tnacrocapsularis  Reuss 
M.  sandrae  Reuss 
M.  bratnae  Reuss 


39; 
40 
41 
42 
43; 
44 
45 
46 
47 
48 
49 
50 
51 
52 
53 
54 

55; 

56 
57 
58; 
59 
6O: 
61 
62 
63 


Genus  2    Ll^SfTOSPDRA    Plehn 
[6  species] 
L.  cerebrdis  (Hofer)  Plehn  (type  species) 
L.  midtiplicata  Reuss 
L.  encephalina  Mulsow 
L.  asymmetrica  Parisi 
L.  acuta  (Fujita)  Kudo 
L.  dermatobia  Ishii 

DAE    Thelohan 

M.  cyprinicola  Reuss 

M.  balleri  Reuss 

M.  squamae  Keysselitz 

M.  cordis  Keysselitz 

M.  musculi  Keysselitz 

M.  sp.  Miyairi 

M.  sp.  Wegener 

M.  permagnus  Wegener 

M.  roiundus  Nemeczek 

M.  minutus  Nemeczek 

M.  sp.  Lebzelter 

M.  magnus  Awerinzew 

M.  carassii  Klokacewa 

M.  sp.  Southwell 

M.  funduli  Kudo 

M.  pleuronectidae  (Hahn) 

M.  capsulatus  Davis 

M.  nodularis  Southwell  et  Prashad    • 

M.  hylae  Johnston  et  Bancroft 

M.  aureatus  Ward 

M.  miyairii  nov.  spec. 

M.  lioi  nov.  spec. 

M.  orbiculatus  nov.  spec. 

M.  discrepans  nov.  spec. 

M.  mesentericus  nov.  spec. 


Genus  2    HENNEGUYA    Th61ohan 
[32  species] 

1)  H.  psorospermica  Thelohan  (type  spe- 

cies) 

2)  H.  texta  (Cohn)  Labb6 

3)  H.  minuta  (Cohn)  Labb6 

4)  H.  oviperda  (Cohn)  Labb6 

5)  H.  lobosa  (Cohn)  Labb6 

6)  H.  peri-intestinalis  C6p6de 

7)  H.  media  Thelohan 

8)  H.  brevis  Th6lohan 

9)  H.  schizura  (Gurley)  Labb6 
10)  H.  creplini  (Gurley)  Labb6 


12 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


t250 


11)  H.  linearis  (Gurley)  Labb6 

12)  H.  gurleyi  Kudo 

13)  H.  strongylura  (Gurley)  Labb6 

14)  H.  monura  (Gurley)  Labb6 

15)  H.  kolesnikovi  (Gurley)  Labb6 

16)  H.  macrura  (Gurley)  Th61ohan 

17)  H.  zschokkei  (Gurley)  Doflein 

18)  H.  sp.  (Gurley)  Labb6 

19)  H.  sp.  (Gurley)  Labbe 

20)  H.  tenuis  Vaney  et  Conte 

21)  H.  nusslini  Schuberg  et  Schroder 

22)  H.  legeri  C6p6de 

23)  H.  acerinae  Schroder 

24)  H.  gigantea  Nemeczek 

25)  E.  (?)  sp.  Nemeczek 

26)  H.  gasterostei  Parisi 

27)  H.  neapolitana  Parisi 

28)  H.  wisconsinensis  Mavor 

29)  E.  brackyura  "Ward 

30)  E.  sdminicola  Ward 


31)  H.  miyairii  nov.  spec. 

32)  E.  mictospora  nov.  spec. 

Genus  3    HOFERELLUS    Berg 
[1  species] 

I)  H.  cyprini  Doflein 

Appendix:  Myxosporidia  of  unknown  genera 
and  species  [1 1  formsl 

1)  Gen.  et  spec,  incert.  Leydig 

2)  Gen.  et  spec,  incert.  Leydig 

3)  Gen.  et  spec,  incert.  Leydig 

4)  Gen.  et  spec,  incert.  Heckel  et  Kner 

5)  Gen.  et  spec,  incert.  Borne 

6)  Gen.  incert.  merlucii  Perugia 

7)  Gen.  incert.  congri  Perugia 

8)  Gen.  et  spec,  incert.  Linton 

9)  Gen.  et  spec,  incert.  Mingazzini 
10)  Gen.  et  spec,  incert.  Nufer 

II)  Gen.  et  spec,  incert.  Mavor 
12)  Gen.  et  spec,  incert.  Mavor 


251]  STUDIES  ON  MYXOSPORIDIA— KUDO  13 

DISTRIBUTION  OF  MYXOSPORIDIA 

A.  GEOGRAPHICAL  DISTRIBUTION 

As  will  be  seen  from  List  III,  Myxosporidia  are  common  parasites  of 
fish  in  various  parts  of  the  world. 

It  is  interesting  to  notice  that  the  same  species  are  found  among  fresh- 
water or  marine  fish  from  waters  in  widely  separated  countries.  It  is 
possible  to  think  that  Myxosporidia  in  marine  fish  may  be  carried  into 
remote  waters  by  the  migration  of  their  hosts,  while  those  infecting  fresh- 
water fish  may  be  brought  from  one  place  to  another  by  the  transportation 
of  infected  fish  for  breeding  purpose,  etc.  It  should  be  noted  in  this  con- 
nection that  no  intermediate  host  has  yet  been  found  in  relation  to  myico- 
sporidiosis. 

The  foUowings  are  the  common  species  found  in  diflferent  localities: 

Leptotheca  parva  Th^l.  Marseille,  Bergen 

Ceratomyxa  sphaerulosa  Th61.  Monaco,  Roscoff,  Bergen 

C.  appendiculata  Th61.  RoscofiE,  Marseille,  Rovigno 

C.  drepanopsetlae  Awerinzew  Murman  coast,  Bergen,  Woods  Hole 

Chloromyxum  leydigi  Ming.  Roscoff,  Monaco,  Napoli,  Rovigno,  Beaufort 

C.  quadratum  Th61.  Roscoff,  Marseille,  Napoli,  Beira 

Sphaerospora  elegans  Th61.  Bretagne,  Karlsruhe,  Lago  di  Garda 

5.  diver  gens  Th61.  Napoli,  Roscoff,  Smalfjorden 

Myxidium  lieberkiihni  Biitsch.  Lago    Maggiore,    France,    Germany,    Lake 

Mendota,  Georgian  Bay 

M.  incurvatum  Th61.  Napoli,  Monaco,  Roscoff,  Bergen,  Beaufort 

M.  bergense  Auerbach.  Bergen,  St.  Andrews 

M.  oviforme  Parisi  Napoli,  Norwegian  coast 

Sphaeromyxa  balbianii  Th61.  Roscoff,  Napoli,  Beaufort 

Myxosoma  dujardini  Th61.  France,  Germany,  Tokio(?) 

On  the  other  hand,  some  species  are  limited  to  certain  localities.  Five 
species  classified  in  the  genus  Sinuolinea  by  Davis  are  reported  only  from 
Beaufort,  N.  C,  U.  S.  A.  The  two  species  of  the  genus  Wardia  have  been 
found  solely  in  the  state  of  Illinois,  U.  S.  A. 

More  detailed  data  are  shown  in  the  following  list. 

LIST  II 

ASIA 
I   Nippon 
Myxosporidia  of  fresh  water  fish 
1)  Northern  Part  (Hokkaido) 

Sapporo :  Mitraspora  cy print  Fujita 
Chloromyxum  koi  Fujita 
Sphaerospora  angulata  Fujita 
Lentospora  acuta  (Fujita)  Kudo 


14 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[252 


2)  Central  part  (Hondo) 

Tokio:    MUraspora  cy print  Fujita 

Chlorotnyxum  misgurni  Kudo 
Chlorotnyxum  fujitai  Kudo 
Spkaerospora  carassii  nov.  spec. 
Myxidiutn  kagayamai  nov.  spec. 
Zsckokkdla  acheilognathi  Kudo 
Myxosoma  dujardini  (?)  Th6lohan 
Myxobolus  toyamai  Kudo 
Myxobolus  misgurni,  nov.  spec. 
Myxoholus  koi  nov.  spec. 
Numazu:  Myxidiutn  anguiUae  Ishii 
Lentospora  dertnatobia  Ishii 

3)  Southern  part  (Kiushiu) 
Fukuoka:  Myxobolus  sp.  Miyairi 

Myxobolus  miyairii  nov.  spec 
Henneguya  miyairii  nov.  spec. 


Katwan,  Mirzapore  (UJP.) 
Mirpur,  Decca  district: 


Bombay: 


n  India 
A.  Myxosporidia  of  fresh-water  fish 
Myxobolus  sp.  Southwell 
Myxobolus  rohitae  Southwell  et  Prashad 
Myxobolus  sent  Southwell  et  Prashad 
Myxobolus  nodularis  Southwell  et  Prashad 

B.  Myxosporidian  of  reptiles 
Myxidium  mackiei  Bosanquet 


m    BUSMA 

In  the  vicinity  of  Ruby  Mines:  Spkaerospora  sp.  Southwell  et  Prashad 

rV   Kamtschatka 
?Henneguya  salminicola  Ward 

AUSTRALIA 
Myxosporidian  of  amphibia 
In  the  vicinity  of  Sidney:  Myxobolus  hylae  Johnston  et  Bancroft 


NUe: 


1)  Indian  Ocean 
Algoa  Bay: 
Beira: 

East  London: 
Lorenfo  Marques: 


AFRICA 

A.  Myxosporidia  of  fresh-water  fish 
Myxobolus  unicapsulatus  Gurley 
Henneguya  strongylura     Gurley 

B.  Myxosporidia  of  marine  fish 


Chioromyxum  magnum  Awerinzew 
Chloromyxum  quadraium  Th61ohan 
Chioromyxum  magnum  Awerinzew 
Ceratomyxa  tylosuri  Awerinzew 
Ceratomyxa  spari  Awerinzew 
Ceratomyxa  sp(?).  Awerinzew 
Ceratomyxa  sp  (?).  Awerinzew 
Sphaeromyxa  exneri  Awerinzew 
2)  South  Atlantic  Ocean 

Luderitz  Bay:         Chioromyxum  magnum  Awerinzew 


253] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


15 


NORTH  AMERICA 


Myxobolus  globosus  Gurley 
Myxobolus  globosus  Gurley 
Myxobolus  sp.  Kudo 
Henneguya  monura  Gurley 


Homer  Park,  HI.: 
Salt  Fork,  Urbana,  HI,: 

Crystal  Lake,  Urbana,  111. 


I   United  States 
A.  Myxosporidia  of  fresh-water  fish 

1)  From  Rivers  emptying  into  Atlantic  Ocean 

Carlius,  Va.  (tribt.  of  Potomac  River) :  Myxobolus  transovalis  Gurley 
Columbia,  S.  C.  (Santee  River) : 
Kinston,  N.  C.  (Neuse  River) : 
West  Falmouth,  Mass. : 
Woodbury,  N.  J.  (Delaware  River) : 

2)  From  Lakes  and  Rivers  opening  into  the  Gulf  of  Mexico 

Fox  River,  trib.  Mbsissippi:  Myxobolus  globosus  Gurley 
Lake  Mendota,  Wis.:  Myxidium  lieberkuhni  Btitschli 

Henneguya  wisconsinensis  Mavor  et  Strasser 
NechesRiver,  Palestin,  Tex.:  Henneguya  macrura  (Gurley)  Th61ohan 
Storm  Lake,  la. :  Henneguya  gurleyi  Kudo 

Stony  Creek,  111. :  Chloromyxum  trijugutn  nov.  spec. 

Myxobolus  orbiculatus  nov.  spec. 

Henneguya  mictospora  nov.  spec. 

Chloromyxum  trijugum  nov.  spec. 

Myxobolus  orbiculatus  nov.  spec. 

Wardia  ovinocua  nov.  gen.  nov.  spec. 

Chloromyxum  catostomi  nov.  spec. 

Myxobolus  discrepans,  nov.  spec. 

Mitraspora  elongate  nov.  spec. 

Myxidium  americanum  nov.  spec. 

Myxobolus  mesentericus  nov.  spec. 

3)  From  the  rivers  opening  into  the  Great  Lakes 

Black  River,  Ohio:  Gen.  et  spec,  incert.  Linton 
Put-In-Bay,  Ohio:      Myxobolus  aureatus  Ward 
Henneguya  brachyura  Ward 

B.  Myxosporidia  of  marine  fish  (Atlantic  Ocean) 

Beaufort,  N.  C. :         Leptotheca  fusiformis  T>a.v\& 
Leptotheca  scissura  Davis 
Leptotheca  lobosa  Davis 
Leptotheca  glomerosa  Davis 
Ceratomyxa  mesospora  Davis 
Ceratomyxa  sphairophora  Davis 
Ceratomyxa  taenia  Davis 
Ceratomyxa  attenuata  Davis 
Ceratomyxa  recurvata  Davis 
Ceratomyxa  lunata  Davis 
Ceratomyxa  abbreviata  Davis 
Ceratomyxa  flagellifera  Davis 
Ceratomyxa  agglomerata  Davfe^  \^^^ 
Ceratomyxa  amorpha  Davis 
Ceratomyxa  m^nospora  Davis 
Ceratomyxa  streptospora  Davis 
Ceratomyxa  aggregata  Davis 
Ceratomyxa  undulata  Davis  > 


16 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[254 


Woods  Hole,  Mass: 


Locality  imrecorded: 
Sycamore,  SL: 


Ceratomyxa  navictdaria  Davis 
Ceratomyxa  spinosa  Davis 
Myxoproteus  cordiformis  Davis 
Myxoproteus  comutus  Davis 
Chloromyxum  leydigi  Mingazzini 
Chloromyxutn  granulosutn  Davis 
Sphaerospora  polymorpha  Davis 
Sinudinea  dimorpha  Davis 
Sinuolinea  capsularis  Davis 
Sinuolinea  arboresqens  Davis 
Sinuolinea  opacita  Davis 
Sinuolinea  brachiophora  Davis 
Myxidium  incurvaium  Th^lohan 
Myxidium  glutinosum  Davis 
Myxidium  phyttium  Davis 
Sphaeromyxa  balbianii  Thfilohan 
Zschokkella  globulosa  Davis 
Myxoboliis  capsulaius  Davis 
Ceratomyxa  drepanopsettae  Awerinzew 
Chloromyxum  funduli  Hahn 
Chloromyxum  dupeidae  Hahn 
Myxosoma  funduli  Kudo 
Myxobdus  lintoni  Gurley 
Myxobolus  funduli  Kudo 
Myxobolus  pleuronectidae  Hahn 
Henneguya  schizura  (Gurley)  LabM 
C.  Myxosporidian  of  Amphibia 
Wardia  ohlmacheri  (Gurley)  Kudo 


II   Canada 
A.  Myxosporidia  of  fresh-water  fish 
Georgian  Bay  (south,  part) :  Myxidium  lieberkuhni  Biitschli 
Myxobolus  notatus  Mavor 

Gen.  et  spec,  incert.  Mavor 

i 

B.  Myxosporidia  of  marine  fish  (Atlantic  Ocean) 
Passamaquoddy  Bay  (at  or 
near  the  mouth  of  St.  Croix 

River),  New  Brunswick:     Ceratomyxa  acadiensis  Mavor 
Myxidium  bergense  Auerbach 
M.  sp.  Mavor 
Gen.  et  spec,  incert.  Mavor 

m  Alaska 
Klutina  Lake:  Chloromyxum  wardi  nov.  spec. 
Stickeen  River:  Henneguya  salminicola  Ward 


SOUTH  AMERICA 
A.  Myxosporidia  of  fresh-water  fish  from  the  waters  connected  with  Atlantic  Ocean 
Guiana:      Myxobolus  inaequalis  Gurley 
Surinam:  Myxobolus  inaequalis  Gurley 
Locality?:  Henneguya  linearis  (Gurley)  Labb6 


255]  STUDIES  ON  MYXOSPORIDIA—KUDO  17 

B.  Myxosporidian  of  marine  fish  (Atlantic  Ocean) 
Rio  de  Janeiro:  Myxidium  striatum  Cunha  et  Fonseca 

C.  Myxosporidian  of  Amphibia 
Brazil:  Spkaeromyxa  immersa  (Lutz)  Th61ohan 

EUROPE 
I  Italy 
A.  Myxosporidia  of  fresh-water  fish  from  lakes  and  rivers  opening  into  Adriatic  Sea 
Lago  di  Como:      MUraspora  caudata  (Parisi)  Kudo 

Myxidium  lieberkiikni  BUtschli 
Lago  di  Garda:     Sphaerospora  degans  Thfilohan 

Henneguya  gasterostei  Parisi 
Lago  di  Varamo :  Henneguya  minuta  (Cohn) 
Lago  Maggiore:    Myxidium  lieberkiikni  Biitschli 
Milano:  Myxidium  lieberkiikni  BUtschli 

Myxobolus  pfeifferi  Thfilohan 
Pa  via:  Myxobolus  gigas  AueThach. 

Myxobolus  ellipsoides  Th^lohan 

Henneguya  peri-intestinalis  C6p6de 
Ticino  River:        Henneguya  minuta  fCohn) 

B.  Myxosporidia  of  marine  fish 

1)  Ligurian  Sea 

Genova :  Chloromyxum  leydigi  Mingazzini 
Gen.  incert.  merluccii  Perugia 
Gen.  incert.  congri  Perugia 

2)  Tyrrhenian  Sea 

Napoli:  Leptotkeca  agilis  Th61ohan 

Leptotkeca  elongata  Thflohan 
Ceratomyxa  arcuata  Th^Iohan 
Ceratomyxa  appendiculata  Th^lohan 
Ceratomyxa  truncata  Thelohan 
Ceratomyxa  inaequalis  Doflein 
Ceratomyxa  linospora  Doflein 
Myxoproteus  ambiguus  (Th61.)  Doflein 
Chloromyxum  leydigi  Mingazzini 
Ckoromyxum  quadraium  Thelohan 
Spkaerospora  diver  gens  Thelohan 
Myxidium  incurvatum  Th61ohan 
Myxidium  giganteum  Doflein 
Myxidium  depressum  Parisi 
Myxidium  oviforme  Parisi 
Spkaeromyxa  balbianii  Thdlohan 
Spkaeromyxa  incurvata  Doflein 
Spkaeromyxa  sabrazesi  Laveran  et  Mesnil 
Lentospora  asymmetrica  Parisi 
Myxobolus  exiguus  Th61ohan 
Myxobolus  miilleri  BUtschli  -^„_^^ 

Henneguya  neapolitana  Parisi 

II  Monaco 
Mjrxosporidia  of  fish  from  Ligurian  Sea 
Leptotkeca  elongata  Th61ohan 
Ceratomyxa  sphaerulosa  Thelohan 


18  ILUNOJS  BIOLOGICAL  MONOGRAPHS  [256 

Ceratomyxa  arcuata  Th61oban 
Ceratotnyota  pallida  Th61ohan 
Ceratomyxa  herouardi  Georg^vitch 
Ceratomyxa  sp.  Georg6vitch 
Chloromyxum  leydigi  Mingazzini 
Myxidium  incurvatum  Th61ohan 
Sphaeromyxa  sabrazesi  Laveran  et  Mesnil 

in  France 
A.  Myxosporidia  of  fresh-water  fish 

1)  From  Rivers  opening  into  Atlantic  Ocean 

Aigne:  Myxobolus  pfeiferi  Th61ohan 

Bretagne:  Sphaerospora  elegans  Th61ohan 

Lorraine:  Myxobolus  ovtformis  Th61ohan 

Nancy:  Myxobolus  pjeiferi  Thelohan 

Mame:  Myxobolus  pfeifferi  Th61ohan 

Seine:  Myxobolus  pfeijfferi  Th61ohan 

Paris:  Chloromyxum  fluviatUe  Thilohan 

Wimereux:  Myxidium  giardi  C6p^de 

2)  From  Rivers  opening  into  Mediterranean  Sea 

Dauphin6:  Myxobolus  miilleri  Butschli 

Drac  River:         Myxobolus  miilleri  Butschli 

Myxobolus  pfeiferi  Thelohan 
Grenoble:  Chloromyxum  cristatum  L^ger 

Is&re  River:  Myxidium  barbatulae  C6pSde 

Myxobolus  oviformis  Thelohan 
Myxobolus  miilleri  Butschli 
Myxobolus  cycloides  Gurley 
Henneguya  Ugeri  C6pede 
Lac  d'Annecy:     Myxobolus  miilleri  Biitschli 
Lac  de  Paladru:  Myxobolus  cycloides  Gurley 
Lac  du  Bourget:  Myxobolus  obesus  Gurley  • 

Henneguya  peri-intestinalis  C6p6de 
Lyon?:  Henneguya  tenuis  Vaney  et  Conte 

Rhdne  River:       Myxobolus  pfeiferi  Th61ohan 
Sa6ne  River:        Myxobolus  pfeiferi  Th61ohan 

B.  Myxosporidia  of  marine  fish 
1)  From  Atlantic  Ocean 

Arcachon  Sphaeromyxa  sabrazesi  Laveran  et  Mesnil 

Concameau:  Ceratomyxa  arcuata  Th61ohan 

Chloromyxum  leydigi  Mingazzini 

Chloromyxum  quadratum  Th61ohan 

Sphaerospora  divergens  Th61ohan 

Myxidium  incurvatum  Thelohan 

Sphaeromyxa  balbianii  Thelohan 
Le  Croisic:  Leptotheca  dongata  Thelohan 

Leptotheca  parva  Thelohan 

Leptotheca  renicola  Thelohan 

Ceratomyxa  appendiculata  Th61ohan 

Myxoproteus  ambiguus  (Th61.)  Doflein 

Sphaerospora  rostrata  Thelohan 


257] 


STUDIES  ON  MYXOSPORIDJA—KUDO 


19 


CoDcameau:  Ceratomyxa  arcuataThiloha-n 

Chloromyxum  leydigi  Mingazzini 
CUoromyxum  quadratum  Thfilohan 
Sphaerospora  divergens  Thdlohan 
Myxidium  incurvatum  Th61ohan 
Sphaeromyxa  balbianii  Th61ohan 

Roscoff:  Cer atomy xa  sphaertdosa  Th^lohan 

Ceratomyxa  arcuata  Th6Iohan 
Ceratomyxa  appendiculata  Th61ohan 
Chloromyxum  leydigi  Mingazzini 
Chloromyxum  quadratum  Th61ohan 
Sphaerospora  rostrata  Thdlohan 
Sphaerospora  divergens  Th61ohan 
Myxidium  incurvatum  Thdlohan 
Myxidium  gadi  Georg^vitch 
Sphaeromyxa  balbianii  Th61ohan 
Sphaeromyxa  sabrazesi  Laveran  et  Mesnil 
Sphaeromyxa  gasterostei  Georg6vitch 
Myxobolus  miilleri  Biitschli 

Le-Vivier-sur-mer:   Leptotheca  parva  Thdlohan 

Myxidium  sphaericum  Thdlohan 
Myxobolus  exiguus  Th^lohan 

St.-Valery-en-caux:  Ceratomyxa  sphaerulosa  Th6Iohan 

2)  From  Mediterranean  coast 

Marseille:  Leptotheca  elongata  Th^lohan 

Leptotheca  parva  Th^lohan 
Leptotheca  renicola  TWlohan 
Leptotheca  hepseti  Thelohan 
Ceratomyxa  arcuata  Thelohan 
Ceratomyxa  pallida  Thelohan 
Ceratomyxa  globulifera  Th6Iohan 
Ceratomyxa  appendiculata  Th61ohan 
Ceratomyxa  reticularis  Th61ohan 
Chloromyxum  leydigi  Mingazzini 
Sphaerospora  rostrata  Thelohan 
Myxidium  incurvatum  Th61ohan 
Myxidium  sphaericum  Thelohan 
Sphaeromyxa  balbianii  Th61ohan 
Myxobolus  exiguus  Thelohan 

Banyuls:  Leptotheca  elongata  Thelohan 

Leptotheca  polymorpha  (Th61.)  Labb6 
Ceratomyxa  arcuata  Thelohan 
Ceratomyxa  globulifera  Thelohan 
Ceratomyxa  appendiculata  Thelohan 
Ceratomyxa  reticularis  Th61ohan 
Chloromyxum  leydigi  Mingazzini 
Sphaerospora  rostrata  Thelohan 
Myxidium  incurvatum  Thelohan 
Myxidium  sphaericum  Thelohan 
Sphaeromyxa  balbianii  Thelohan 
Myxobolus  exiguus  Thelohan 


20 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


1258 


Villefranche:  Ceratomyxa  pallida  Thfilohan 

Ceratotnyxa  truncata  Th61ohan 
Ceratomyxa  coris  Georg6vitch 
Sphaeromyxa  balbianii  Th61ohan 

Locality  unknown:  Leptotheca  agilis  Th61ohan 

Leptotheca  perlata  (Gurley)  Labb6 
Myxidium  lieberkUhni  Biitschli 
Myxidiutn  histophilutn  Th61ohan 
Myxosoma  dujardini  Th61ohan 
Myxobolus  piriformis  Th^lohan 
Myxobolus  dispar  Th61ohan 
Myxobolus  obesus  TWlohan 
Henneguya  psorospermica  Thelohan 
Henneguya  media  Thelohan 
Henneguya  brevis  Thelohan 
Hoferellus  cyprini  Doflein 

C.  Myxosporidian  in  a  reptile 
Myxidium  danilewskyi  Laveran 

IV  Germany 
A.  Myxosporidia  of  fresh-water  fish 
1)  From  Rivers  opening  into  North  Sea 

Throughout  country:  Myxobolus  cyprini  Doflein 


Berlin: 
Bodensee: 

Gutach: 

Karlsruhe  and  its 
vicinity: 


Leipzig: 
Mosel: 


Neckar: 


Rhine: 


Henneguya  ovipcrda  (Cohn) 
Chloromyxum  dubium  Auerbach 
Myxobolus  mulleri  Biitschli 
Myxobolus  neurobitis  Schuberg  et  Schroder 
Henneguya  niisslini  Schub.  et  Schroder 
Chloromyxum  mucronatum  Gurley 
Sphaerospora  elegans  Thdlohan 
Myxidium  lieberkUhni  Biitschli 
Myxidium  pfeijfferi  Auerbach 
Myxidium  macrocapstdare  Auerbach 
Henneguya  oviperda  (Cohn) 
Henneguya  lobosa  (Cohn) 
Myxobolus  gigas  Auerbach 
Myxobolus  sp.  Gurley 
Myxobolus  pfeiferi  Thelohan 
Myxobolus  squamae  KeysseUtz 
Myxobolus  cordis  KeysseUtz 
Myxobolus  musculi  KeysseUtz 
Myxobolus  exiguus  Thelohan  (Heidelberg) 
Myxobolus  mulleri  ButschU 
Myxobolus  pfeiferi  Th61ohan 
Myxobolus  squamae  KeysseUtz 
Myxobolus  cordis  KeysseUtz 
Myxobolus  musculi  KeysseUtz 
Henneguya  psorospermica  Th6lohan 
Henneguya  acerinae  Schroder  (Heidelberg) 
Myxidium  lieberkUhni  Btitschli 
Myxobolus  mulleri  ButschU 
Henneguya  psorospermica  Th61ohan 
Lentospora  encephalina  Mulsow 


259] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


21 


2)  From  Rivers  opening  into  Baltic  Sea 


AUe: 
Frisches  Haff: 


Kurisches  Haff : 


Masurische  Seen: 


Pregel: 


Weichsel: 
3)  Localities  unknown: 


Myxobolus  midleri  BUtschli 
Myxidium  lieberkuhni  Butschli 
Myxosoma  dujardini  Th61ohan 
Myxobolus  piriformis  Thdlohan 
Myxobolus  dispar  Th61ohan 
Myxobolus  exiguus  Thdlohan 
Myxobolus  ovifortnis  Th61ohan 
Myxobolus  tniilleri  Butschli 
Myxobolus  cycloides  Gurley 
Myxobolus  anurus  Cohn 
Myxobolus  sp.  Wegener 
Myxobolus  permagnus  Wegener 
Henneguya  psorospermica  Thelohan 
Henneguya  texta  (Cohn) 
Henneguya  minuta  (Cohn) 
Henneguya  lobosa  (Cohn) 
Henneguya  crepUni  (Gurley) 
Myxosoma  dujardini  Thelohan 
Myxobolus  exiguus  Thelohan 
Myxobolus  ovifortnis  Thelohan 
Myxobolus  cycloides  Gurley 
Henneguya  psorospermica  Th61ohan 
Henneguya  texla  (Cohn) 
Henneguya  creplini  (Gurley)  Labb6 
Sphaerospora  masovica  Cohn 
Myxobolus  dispar  Th61ohan 
Myxobolus  ellipsoides  Thelohan 
Myxobolus  cycloides  Gurley 
Myxobolus  anurus  Cohn 
Henneguya  psorospermica  Th61ohan 
Henneguya  texta  (Cohn) 
Myxosoma  dujardini  Thelohan 
Myxobolus  piriformis  Thelohan 
Myxobolus  dispar  Thelohan 
Myxobolus  exiguus  Thelohan 
Myxobolus  oviformis  Thdlohan 
Myxobolus  miilleri  Butschli 
Myxobolus  cycloides  Gurley 
Myxobolus  anurus  Cohn 
Myxobolus  permagnus  Wegener 
Henneguya  psorospermica  Thelohan 
Henneguya  texta  (Cohn) 
Henneguya  minuta  (Cohn) 
Henneguya  lobosa  (Cohn) 
Henneguya  creplini  (Gurley)  Labb6 
Myxobolus  cyprini  DoSein 

Chloromyxum  leydigi  Mingazzini 
Myxidium  lieberkiihni  Butschli 
Myxidium  sp.  Gurley 
Lentospora  cerebralis  (Hofer)  Plehn 


22 


ILUNOIS  BIOLOGICAL  MONOGRAPHS 


(260 


Helder: 


Henneguya  sckizura  (Gurley)  Labb6 
HojereUus  cyprini  Doflein 
Gen.  et  spec,  incert.  Leydig 
Gen.  et  spec,  incert.  Leydig 
Gen.  et  spec,  incert.  Leydig 
Gen.  et  spec,  incert.  Bome 

V  Netheuland 
Myxosporidian  of  marine  fish 
CMoromyxum  quadratum  Th61ohan 


VI  England 
M3rxosporidia  of  marine  fish 
Firth  of  Qyde,  More- 

camb,  etc.:  Myxobolus  aeglefini  Auerbach 

Liverpool  (?) :  Sphaerospora  platessae  Woodcock 


Abelvaer: 


Bergen: 


Bergsfjord: 

Boadsfjord: 
Bodd: 

Finkongkjeilen: 

GronSy: 

Hammerfest: 

Harstad: 

Honnigsvaag: 

Kabelvaag: 


Vn  Norway 
Mjrxosporidia  of  marine  fish 
Myxidium  bergense  Auerbach 
Myxidium  ovijorme  Parisi 
Zschokkelia  hildae  Auerbach 
Myxobolus  aeglefini  Auerbach 
Leptotheca  parva  Th^lohan 
Leptotheca  macrospora  Auerbach 
Leptotheca  informis  Auerbach 
Leptotheca  longipes  Auerbach 
Ceratomyxa  sphaerulosa  Thelohan 
Myxidium  incurvatum  Thfilohan 
Myxidium  inflatum  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  procerum  Auerbach 
Sphaeromyxa  heUandi  Auerbach 
Zschokkelia  hildae  Auerbach 
Myxobolus  aeglefini  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Zschokkelia  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  ovijorme  Parisi 
Zschokkelia  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Myocidium  bergense  Auerbach 
Zschokkelia  hildae  Auerbach 
Ceratomyxa  drepanopsettae  Awerinzew 


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STUDIES  ON  MYXOSPORIDIA—KUDO 


23 


Myxidium  bergense  Auerbach 
Zschokkella  hildae  Auerbach 
Zschokkdla  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  bergense  Auerbach 
LepMheca  parva  Thdlohan 
Lepiotheca  macrospora  Auerbach 
Myxidium  oviforme  Parisi 
Zschokkella  hildae  Auerbach 
Myxobolus  aeglefini  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkella  hildae  Auerbach 
Zschokkella  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkella  hildae  Auerbach 
Ceratomyxa  drepanopsettae  Awermzew 
Myxidium  bergense  Auerbach 
Myxidium  oviform^  Parisi 
Zschokkella  hildae  Auerbach 
Zschokkella  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Sphaerospora  divergens  Thdlohan 
Zschokkella  hildae  Auerbach 
Leptotheca  parva  Thelohan 
Myxidium  bergense  Auerbach 
Myxidium  bergense  Auerbach 
Zschokkella  hildae  Auerbach 
Leptotheca  informis  Th61ohan 
Ceratomyxa  drepanopsettae  Awerinzew 
Myxidium  bergense  Auerbach 
Sphaeromyxa  hellandi  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  oviforme  Parisi 
Zschokkella  hildae  Auerbach 
Zschokkella  hildae  Auerbach 
Myxidium  bergense  Auerbach 
Myxidium  oviforme  Parisi 
Zschokkella  hildae  Auerbach 
Myxobolus  aeglefini  Auerbach 


VIII  Switzerland 
Myxosporidia  of  fresh-water  fish 
1)  From  Lakes  connected  with  North  Sea 

Neuchatel:  Myxobolus  fuhrmanni  Auerbach 

Myxobolus  miUleri  Biitschli 
Henneguya  oviperda  (Cohn) 
Henneguya  zschokkei  (Gurley)  Doflein 
Thun:  Henneguya  zschokkei  (Gurley)  Doflein 

Zurich:  Henneguya  zschokkei  (Gurley)  Doflein 

Lucerne:  Myxosoma  dujardini  Th61ohan 

Myxobolus  ellipsoides  Thelohan 


Kiberg: 
Kirkenes: 
Eliistiana: 
Kristiansand: 


Lodingen: 
Makur: 

Mosjoen: 

Nusfjord: 

RSrvik: 

Rossfjord: 

SkjervS: 

Skjotningsberg: 

Smalfjorden: 

Stavanger: 

Svolvaer: 

TjamS: 


Torghatten: 
Trondhjem: 


Vikholmen: 
Vardo: 


24 


ILUNOJS  BIOLOGICAL  MONOGRAPHS 


[262 


Myxobolus  ovifortnis  Th61ohan 
Myxobolus  midleri  Biitschli 
Henneguya  psorospermka  Th^lohan 
Henneguya  texta  (Cohn) 
Henneguya  zschokkei  (Gurley)  Doflein 
Gen.  et  spec,  incert.  Nufer 

Wallen:  Henneguya  zschokkei  (Gurley)  Doflein 

2)  From  Lake  connected  with  Mediterranean  Sea 

Geneva:  Myxobolus  sphaeralis  Gurley 

Henneguya  zschokkei  (Gurley)  Doflein 

IX  AtJSTKIA 

A.  Myxosporidia  of  fresh-water  fish 

1)  From  Wvers  opening  into  Black  Sea 

Danube  tributaries     Chloromyxum  thynwlli  Lebzelter 
and  Neusiedler:        Myxosotna  (?)  lobatum  Nemeczek 
Myxobolus  aeglefini  Auerbach 
Myxobolus  cyprini  Doflein 
Myxobolus  rotundus  Nemeczek 
Myxobolus  minutus  Nemeczek 
Myxobolus  sp.  Lebzelter 
Henneguya  acerinae  Schroder 
Henneguya  gigantea  Nemeczek 

2)  From  Rivers  opening  into  North  Sea 


Prag: 


Krakau: 


Rovigno: 


Locality  unknown: 


Vienna: 


Pergrad  (Danube): 


Volga  (to  Caspian 
Sea): 


Myxosoma  dujardini  Th61ohan 
Myxobolus  ellipsoides  Th61ohan 
Myxobolus  oculi-leucisci  Trojan 
Myxobolus  cyprini  Doflein 
Myxosporidia  of  marine  fish  (Adriatic  Sea) 
Leptotheca  agilis  Thelohan 
Ceratontyxa  pallida  Th61ohan 
Ceralomyxa  appendiculata  Th61ohan 
Myxoproteus  ambiguus  (Th^l.)  Doflein 
Chloromyxum  leydigi  Mingazzini 
Sphaeromyxa  sabrazesi  Laveran  et  Mesnil 
Gen.  et  spec,  incert.  Heckel  et  Kner 
C.  Myxosporidian  of  Amphibia 
Chloromyxum  protei  Joseph 

X  Serbia 
Henneguya  gigantea  Nemeczek 

XI  Russia 

A.  Myxosporidia  of  fresh-water  fish 

Lentospora  multiplicata  Reuss 
Myxobolus  volgensis  Reuss 
Myxobolus  scardinii  Reuss 
Myxobolus  physophilus  Reuss 
Myxobolus  macrocapsularis  Reuss 
Myxobolus  sandrae  Reuss 
Myxobolus  bramae  Reuss 
Myxobolus  cyprinicola  Reuss 
Myxobolus  balleri  Reuss 


263] 


Don  (to  Black  Sea) : 
Locality  unknown : 


STUDIES  ON  MYXOSPORJDI A—KUDO 

Myxobolus  sp.  Gurley 
Zschokkella  nova  Klokacewa 
Myxobolus  tnagnus  Awerinzew 
Myxobolus  carassii  Klokacewa 
Henneguya  kolesnikovi  (Gurley)  Labb6 


25 


B.  Myxosporidia  of  marine  fish  from  Arctic  Ocean 
Murman  coast:  Ceratomyxa  ramosa  Awerinzew 

Ceratomyxa  drepatwpsettae  Awerinzew 
Myxidium  sp.  Awerinzew 
Leptotheca  sp.  Awerinzew 
Chloromyxum  sp.  Awerinzew 

B.  DISTRIBUTION  OF  MYXOSPORIDIA  IN  ANIMALS 

The  number  of  host  species  that  harbor  Myxosporidia  is  237,  as  will  be 
seen  from  List  III. 

Tho  two  incompletely  studied  forms  are  found  in  Annelida  and  Insecta, 
Myxosporidia  are  the  parasites  of  Vertebrata,  especially  of  Pisces,  only  few 
being  found  infecting  Amphibia  and  Reptilia.  They  are  distributed  among 
these  groups  of  animals  as  follows: 

Number  of  host  species 

Annelida 1 

Insecta 1 

Pisces 223 

Amphibia 8 

Reptilia 4 

Gurley  (1894:101-105),  Wasielewsky  (1896:132-148),  Labbe  (1899: 
133-161)  and  Auerbach  (1910:36-45;  1911:471-494)  gave  lists  in  which 
they  recorded  the  names  of  host  species.  Wasielewsky  arranged  the  names 
alphabetically  while  others  listed  them  according  to  their  systematic  order. 
In  the  following  pages,  the  writer  followed  Wasielewsky,  i.e.,  the  names  of 
the  host  species  are  arranged  alphabetically  as  is  supposed  to  be  more 
convenient  in  referring  to  the  host  than  any  form  presented  otherwise. 

LIST  III.     LIST  OF  HOST  SPECIES 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Annelida 
Nais  lacustris  {N.  probo- 
scidea) 

Unknown 

Abdominal  cav- 
ity 

Branchiae 

Myxobolus  sp. 

Chloromyxum  diploxys 

Myxobolus  balleri 
bramae 

Germany 

Insecta 

Tortrix  viridana  L.  (imago) 

Pisces 

Abramis  batterus  L 

France 
Russia 

A.  brama  L     

« 

26 


ILUNOIS  BIOLOGICAL  MONOGRAPHS 


[264 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Branchiae 

Myxobolus  cycloides 

Germany 

i< 

ellipsoides 

"(?) 

« 

exiguus 

France 

(1 

ovifonnis 

France 

(C 

rotundus 

Austria 

Gall-bladder 

Sphaerospora  masovica 

Germany 

Kidney 

Myxobolus  cyprini 

« 
t 

Hungary 

Subcut.    conn. 

gigas 

Germany, 

tissue  of  oper- 

Italy 

culum 

A.  vimba  L 

Branchiae 

cycloides 

Germany 

« 

ellipsoides 

"(?) 

« 

oviformis 

Germany 

Acanthias  acantkias  L 

Gall-bladder 

Chloromyxum  leydigi 

France 

A.  blainvillei 

(1 

magnum 
Henneguya  acerinae 

Africa 

Acerina  cernua  L 

Branchiae 

Germany 

"   .Muscle 

crepiini 

(( 

Conn,  tissue  of 

tenuis 

France 

aliment,  canal 

Eye 

Myxobolus  magnus 

Russia 

Muscle 

Leptotheca  perlata 

France? 

Acheilognathus  lanceolatum 

Gall-bladder, 

Temm.  et  Schl 

Gall-duct 

Zschokkella  acheilognathi 
Myxobolus  cycloides 

Nippon 
Germany 

Alburnus  alburnus  L 

Branchiae 

(A.  lucidus  Heck) 

(( 

dispar 

« 

« 

ellipsoides 

Germany  ? 

It 

oviformis 

France 

"  ,  kidney 

obesus 

(1 

Muscle  and 

dispar 

f( 

spleen 

Eye 

miilleri 

Switzerland 

Alosafinta  Cuv.  var.  lacus- 

tris  Fatio 

Kidney 

Base  of  spines  of 
2nd  dorsal  fin 

Mitraspora  caudata 

Italy 

Ameiurus  tnelas  Raf 

Henneguya  gurleyi 

U.  S.  A. 

Ancyhpsetta  quadrocdlata 

GUI 

Gall-bladder 

Ceratomyxa  undulata 
Myxidium  anguiUae 

« 

AnguUla  japonica  Temm. 

Integument 

Nippon 

etSchL 

« 

Lentospora  dermatobia 
Myxidium  giardi 

« 

A.  vulgaris  Flemm 

Kidney 
Subcutaneous 

France 

Apkredoderus  sayanus  Gill. 

intermusc.  tiss. 

Henneguya  monura 

U.  S.  A. 

Apogon  rex-mtdlorum  Cuv. 

GaU-bladder 

Myxidium  oviforme 

Italy 

Argentina  situs  Nilss 

« 

procerum 
Chloromyxum  quadratum 

Norway 

Ariodes  polystaphylodon 

Muscle 

Africa 

Aspius  rapax  Ag 

Branchiae 

Myxosoma{7)  lobatum 
Myxobolus  miilleri 
Henneguya  acerinae 

Austria 

Aspro  asper  L 

(( 

France 

A.  zingd  Cuv 

« 

« 

2651 


STUDIES  ON  MYXOSPORIDIA—KUDO 


27 


Host 

Organ  Infected 

Myxosporidian 

LocaUty 

Atherina  hepsetus  L 

Gall-bladder 

Leptotheca  hepseH 
Myxoproteus  cornutus 
Myxidium  striatum 

France 

Bairdiella  chrysura 

Urin.  bladder 

U.  S.  A. 

B.  rouchus  Cuv.  et  Val 

GaU-bladder 

Brazil 

Barbus  barbus  L 

Kidney,  spleen, 
intestine,  ovary, 

(5.  fluviatUis) 

Myxobolus  pfeifferi 

France, 

etc. 

Germany 

Inner  surface  of 

squamae 

Germany 

scale 

Muscle  of  ven- 

cordis 

<f 

tricle 

Muscle,  kidney 

muscuH 

« 

B.  plebejus  Val 

(( 

pfeifferi 
miiUeri 

Italy 

Germany 

Burma 

B.  vulgaris  Flem 

Branchiae 

Barilius  barnc ;.. 

Under  scales 

Sphaerospora  sp. 
Myxidium  sphaericum 
Myxidium  sphaericum 
Sphaeromyxa  incurvata 
Chloromyxum  quadratum 
Myxidium  incurvatum 
Sphaerospora  divergens 

Belone  acus  Risso 

GaU-bladder 

it 

France 

B.  belone  L 

« 

Blennius  ocellatus 

« 

Italy 
« 

B.  gattorugine  Brunn 

Kidney 
Gall-bladder 

B.  pholis  L 

France 

Renal  tubules 

« 

Blicca  bjdrkna  L 

Branchiae 

Myxobolus  cydoides 
ellipsoides 

Germany 
"? 

« 

« 

macrocapsularis 

Russia 

« 

oviformis 

Germany 

Box  boops  L 

Gall-bladder     . 

Ceratomyxa  pallida 

France, 

Italy 

B.  salpa  L 

Gall-bladder 

Ceratomyxa  herouardi 
pallida 

Monaco 

(( 

France, 

Italy 

Kidney 

Henneguya  neapolitana 

Italy 

BrevoorUa  tyrannus  Latr.... 

Muscle 

Chloromyxum  clupeidae 

U.  S.  A. 

Brosmius  brosme  Ascanius.. 

GaU-bladder 

Leptotheca  longipes 

Norway 

« 

Sphaeromyxa  hellandi 

({ 

CaUionymus  lyra  L 

« 

Myxidium  incurvatum 

France, 

Norway 

Muscle 

Chloromyxum  quadratum 

France 

Carassius  auratus  L 

Branchiae 

Lentospora  acuta 
Sphaerospora  angulata 

Nippon 
« 

Kidney 

« 

Mitraspora  cyprini 

<( 

Subcutaneous 

tiss.  of  head 

Henneguya  miyairii 

« 

(C.  carassius  L.) 

Body  cavity 
Branchiae 

Myxobolus  sp. 

dispar 

Germany 
« 

u 

Sphaerospora  carassii 

Nippon 

(C.  vulgaris  L.) 

Body  cavity, 
Uver,  intestine 

Myxobolus  carassii 

Russia 

GaU-bladder 

Zschokkella  nova 

(( 

Carcharhinus  limbatus 

(< 

Chloromyxum  leydigi 

U.S.A. 

C.  sp 

« 

Ceratomyxa  flagellifera 

« 

28 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[266 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Carpiodes  difformis 

Branchiae 

Myxobolus  discrepans 

U.  S.  A. 

Catostomus   comtnersonii 

Lac 

Gall-bladder 

Chloromyxum  catostomi 
Sphaeromyxa  hellandi 

« 

Centronotus  gundlus 

(( 

Norway 

Cepola  rubescens  L 

(( 

balbianii 

France 

Cestracion  tiburo 

« 

Chloromyxum  leydigi 

U.  S.  A. 

(1 

Ceratomyxa  mesospora 

a 

C.  zygaena 

(( 

Ceratomyxa  mesospora 

it 

(( 

recurvata 

it 

« 

Chloromyxum  leydigi 

ti 

(i 

Leptoiheca  fusiformis 

it 

Chaetodipterus  faber 

Gall-bladder 

Ceratomyxa  streptospora 

It 

Urin.  bladder 

Myxoproteus  cordiformis 

It 

Chondrostoma  nasus  L 

Branchiae 

Myxobolus  exiguus 

Germany 

" 

Gen.  et  spec,  incert. 

Switzerland 

Tongue 

Gen.  et  spec,  incert. 

? 

Citharus  linguataGthi 

Gall-bladder 

Myxidium  depressum 

Italy 

Clupea  harengus  Young 

(( 

Ceratomyxa  sphaerulosa 

Norway 

Muscle 

Chloromyxum  dupeidae 

U.  S.  A. 

C.  pilchardus  Walb. 

{Alosa  sardina) 

Gall-bladder 

Ceratomyxa  truncata 
Sphaeromyxa  balbianii 

France,  Italy 
Italy 

it 

Cohitis  barbatula  L 

Kidney 
Urin.  bladder 

Myxidium  barbatulae 
Henneguya  legeri 

France 

« 

C.  fossilis  L 

Branchiae, 

Conger  conger  L. 

kidney,    spleen 

Myxobolus  piriformis 

Germany 

{Leptocepkalus  c.) 

Gall-bladder 

Gen.  incert.  congri 

Italy 

Coregonus  lavaretus  L. 

(C.fera) 

Branchiae 

Henneguya  sp. 
Myxobolus  sphaeralis 

France? 

\^^  'J  ^' "/ 

"  (mucosa) 

Switzerland 

Muscle 

Henneguya  kolesnikovi 

Russia 

« 

zschokkei 

Switzerland 

C.  exiguus  albellus 

"  ,  branchia 

it 

« 

C.  wartmanni  nobilis 

«            « 

tt 

« 

Coris  giofredi  Risso 

Gall-bladder 

Ceratomyxa  coris 
it 

France 

C.julisL 

(( 

tt 

Muscle 

Chloromyxum  quadratum 

ti 

Gall-bladder 

Myxidium  oviforme 

Italy 

CoUus  gobis  L 

Branchiae 

Myxobolus  mulleri 
Myxidium  sp. 
Ceratomyxa  inaequaiis 

France 

C.  scorpitis 

« 

Russia 

Crenilabrus  mediterraneus.. 

" 

Italy 

C.  tnelops  L ' 

Eye 

Myxobolus  miiUeri 

Germany 

J  w 

France 

Gall-bladder 

Ceratomyxa  arcuata 

France 

Kidney 

Sphaerospora  diver  gens 

K 

C.  pavo  Cuv.  et  Val 

Gall-bladder 

Ceratomyxa  inaequaiis 
Lentospora  asymmetrica 

Italy 

Kidney 

Italy 

« 

Sphaerospora  diver  gens 

France,  Italy 

Cydopterus  lutnpus  L 

Gall-bladder 

Myxidium  infiatum 

Norway 

267] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


29 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Cyitoscion  regalis 

Gall-bladder 

Myxidium  glutinosum 

U.  S.  A. 

Urin.     bladder, 

ureters 

Sinuolinea  dimorpha 

« 

CypHnodon  variegatus 

Subcutaneous 

tissue 

Myxobolus  lintoni 

(( 

Visceral  conn. 

tissue 

capsulatus 

« 

CypHnus  carpioli 

Branchiae 

cyprinicola 

Russia 

(( 

dispar 

France, 
Germany 

<( 

oviformis 

France  ? 

<c 

toyamai 

Nippon 

tl 

Myxosoma  dujardini 

France 

(( 

Myxobolus  koi 

Nippon 

Brain 

Lentospora  encephalina 

Germany 

Gall-bladder 

Chloromyxum  koi 

Nippon 

Kidney 

Hoferellus  cyprini 

Germany, 
France 

"     liver,  spl. 

Myxobolus  cyprini 

Germany, 
Himgary 

Kidney 

Milraspora  cyprini 

Nippon 

« 

Spkaerospora  angulata 

« 

C.  (Rasbora)  daniconius 

Subcutaneous 

Day 

intermuscular 

tissue 

Myxobolus  sp. 

India 

Muscles 

Myxobolus  nodularis 

India 

Dasybatis  hastatus 

Gall-bladder 

Chloromyxum  leydigi 
Leptotheca  scissura 

U.  S.  A. 

(( 

« 

D.  sabina 

« 

Chloromyxum  leydigi 

« 

Drepanopsetta  platessoides 

Fabr 

(1 

Ceratomyxa  drepanopsettae 

Russia 

Erimyzon  sucetta  oblongus 

Lac  {Catostomus 

Branchiae 

Myxobolus  globosus 

U.  S.  A. 

tuberctdatus) 

Integument 

oblongus 

i( 

Esox  lucius  L.... 

Branchiae 

anurus 

Germany 

(( 

Henneguya  lobosa 

« 

<( 

Henneguya  psorospermica 

France, 
Germany 

Intestinal  wall 

Henneguya  peri-inkstinalis 

France, 
Italy 

Eye  muscle,  etc. 

Henneguya  sckizura 

U.  S.  A. 

Ovary 

Henneguya  oviperda 

Germany, 
Switzerland 

Urin.  bladder 

Myxidium  lieberkUhni 

France,  Ita- 
ly, Canada, 
U.  S.  A., 
Germany 

Fundidus  sp 

Muscle 

Chloromyxum  funduli 

U.  S.  A. 

30 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[268 


Host 


Organ  Infected 


Myxosporidian 


Locality 


P.  diaphanus.. 
F.  keteroditus.. 


F.  majalis.. 


Gadus  aeglefinis  L., 
G.  caUarias  L 


G.  tsmarkii  Nilss.. 


G.merlangus'L.. 


G.  morrhuaJj.... 
G.  poUackius  L.. 
G.  virens  L 


Galeocerda  tigrinus 

Galois  galeus  L.  (G.  canis). 
Gambusia  affinis 


GasterosUus  aculeatus. 


G.  pungitius  L.. 


G.  spinachia 

Gobio  gobio'L. 
(G.  fiuviatUis). 


Gobius  fluviatUis  L 

Gobius  paganeUus  L 

Hdiases  ckromis  Gthr.. 


Hippocampus  brevirostris 
Cuv. 


Muscle 

Branchiae, 

Muscle 

Branchiae 
« 

"  ,    muscle 
Gall-bladder 
Cartilage 
Urin.  bladder 
Cartilage 
Gall-bladder 
Urin.  bladder 
Cartilage,  bone 

of  cranium, 

eye 

Gall-bladder 

Cartilage 

Cartilage 

Gall-bladder 
« 

Urin.  bladder 
Gall-bladder 


Kidney  (r.  t.), 
ovary 


Kidney 
"  (r.  t),  ovary 


Gall-bladder 

Fin 

Fin,  spleen, 
kidney,  liver 
Branchiae 
Body-cavity 
Gall-bladder 


Myxoholus  funduli 

Myxobolus  funduli 
Myxosoma  funduli 
Myxosoma  funduli 
Myxobolus  funduli 
Myxidium  incurvatum 
Myxobolus  aeglefini 
Zschokkella  kildae 
Myxobolus  aeglefini 
Myxidium  oviforme 
Zschokkella  hUdae 


Myxobolus  aeglefini 

Leptotheca  informis 
Myxobolus  aeglefini 
Myxobolus  aeglefini 
Myxidium  gadi 

bergense 
Zschokkella  hildae 
Ceratomyxa  lunata 

sphaerulosa 
Myxidium  incurvatum 

phyUium 

Henneguya  media 

brevis 
Sphaerospora  elegans 
Henneguya  gasterostei 

brevis 

media 
Sphaerospora  elegans 
Sphaeromyxa  gasterostei 

Myxobolus  miilleri 

oviformis 

cycloides 

Gen.  et.  spec,  incert. 

Ceratomyxa  arcuata 

Ceratomyxa  arcuata 


Myxidium  incurvatum 
Sphaeromyxa  sabrazesi 


U.S.  A. 


Germany 

Norway 

Germany 

Norway 

Norway 


Germany, 

England 

Norway 

Germany 

Germany 

France 

Norway 
« 

U.  S.  A. 
France 
U.  S.  A. 


France 

« 

"  ,  Italy 
Italy 
France 


Germany 

France 

Germany 
(( 

Italy 
Italy, 
Monaco 

Italy 
France, 

Italy, 

Himgary 


269] 


STUDIES  ON  MYXOSPORIDJA—KUDO 


31 


Host 

Organ  Infected 

Myxosf>oridian 

Locality 

H.  guUulatus  Cuv 

Gall-bladder 

Sphaeromyxa  sabrazesi 

Fr.,  Hun., 

Hippoglossoides    litnan- 

Monaco 

doides 

Urin.  bladder 

Sphaerospora  diver  gens 

Norway 

Hippoglossus  vulgaris 

Flenun 

Gall-bladder 

Ceratomyxa  drepanopsettae 

Norway 
Russia 

(( 

ramosa 

Hybognathus  nuchalis  Ag.... 

Conn,  tissue  of 

the  head 

Henneguya  macrura 

U.  S.  A. 

Idus  tndanotus  Jleck 

Branchiae 

Myxobolus  eUipsoides 

Germany? 

Muscle 

Leniospora  multiplicata 

Russia 

Labeo  rohita 

Branchiae 

Myxobolus  rohilae 

India 

Fins 

Myxobolus  seni 

« 

Ldbeo  niloticus  Forsk 

? 

Myxobolus  unicapsulatus 

Egypt 

Lahrus  turdus 

GaU-bladder 

Ceratomyxa  linospora 

Italy 
U.  S.  A. 

Leiostomus  xanthurus 

i< 

aggregata 

Lepisosleus  platyslomiis 

Urinary  bladder 

Sphaerospora  sp. 

« 

Lepomts  cyanellus  Raf 

Mesentery 

Myxobolus  mesentericus 

« 

Urin.  bladder 

Henneguya  mictospora 

M 

Kidney 

Mitraspora  elongata 

<( 

L.  humilis  Girard 

Ovary 

Wardia  ovinocua 

l< 

Urinary  bladd. 

Henneguya  mictospora 

« 

L.  tnegalotis  Raf 

Gall-bladder 

Chloromyxum  trijugum 

« 

Leuciscus  cephdus  {Squalis 

Air-bladder, 

France, 

cephalus) , 

branchiae 

Myxobolus  miilleri 

Germany 

Branchiae 

ellipsoides 

«  « 

» 

Gall-bladder 

Chloromyxum  fluviatile 

France 

L.  lucius  L 

Branchiae 

Myxobolus  sp. 

Germany? 

L.  phoxinus  L.  {Phoxinus 

Conn.  tiss.  of 

laevis  Ag.) 

kidney; ovary 

Myxidium  histophilum 
Myxobolus  miilleri 

France 

<< 

« 

L.  rutilus  L 

Branchiae 

ellipsoides 

Germany? 

<i 

miilleri 

c< 

Myxosoma  dujardini 

France 

Conn.  tiss.  un- 

der the  mouth 

muc.  mem- 

brane 

Myxobolus  fukrmanni 

Switzerland 

Opercle,  pseu- 

do  branchiae, 

France, 

kidney 

cydoides 

Germany 

Vitreous  body 

of  eye 

oculi-leucisci 

Austria 

? 

Henneguya  sp. 

Germany 

Heart 

Gen.  et  spec,  incert. 

(( 

Leuciscus  sp 

Branchiae 

Myxobolus  minutus 

Austria 

<( 

Myxosoma  (?)  lobatum 

« 

Lophius  budegassa  Spin 

GaU-bladder 

Ceratomyxa  appendiculata 

Italy, 
France 

L.  piscatorius  Li 

(( 

Ceratomyxa  appendiculata 

<i 

32 


ILUNOJS  BIOLOGICAL  MONOGRAPHS 


(270 


Host 

Organ  Infected 

Mjrxosporidian 

Locality 

L.  piscatorius  L 

Urin.  bladder 

Myxoproteus  atnbiguus 

France, 

Italy, 

' 

Hungary 

Lota  lota  L.  (L.  vulgaris) 

Branchiae 

Myxobdus  midUri 

Germany 

« 

Myxoholus  cydoides 

Germany 

Gall-bladder 

CUoromyxum  dubium 

"  ,  Austria 

Urin.  bladder, 

kidney 

mucronatum 

France 

Urin.  bladder 

Myxidium  lieberkiihni 

France, 
Germany 

(( 

Sphaerospora  degans 

Germany 

Lucioperca  lucioperca  L 

Branchiae 

Henneguya  acerinae 

"  ,    Austria 

(L.  Sandra  Cuv.) 

"     ,  head,  fin, 

circle 

Myxoholus  sp. 

? 

<i 

Henneguya  gigantea 

Aus.,  Servia 

« 

Gen.  et  sp.  incert. 

Austria 

Muscle 

Myxoholus  sandrae 

Russia 

L.  vdgensis  Pall 

Branchiae,  cor- 

nea, dorsal  fin 

volgensis 

« 

Mdanogrammus  aeglefinis.. 

GaU-bladder 

Myxidium  hergense 

Canada 

Menticirrkus  americanus  L. 

<( 

striatum 

BrazU 

Merluccius  merluccius  L. 

(M. vulgaris) 

« 

Ceratomyxa  glohulifera 
Leptotheca  dongata 

France 

V'***  •  ■'^^^i*' »*•/•  ••••••••••••••••••• 

« 

"    ,  Italy 

i< 

Gen.  incert.  merluccii 

Italy 

Micropogon  undulatus 

« 

Ceratomyxa  aggregata 

U.S.A. 

Micropterus  salmoides  Lac . 

Urin.  Bladder 

Henneguya  mictospora 

U.  S.  A. 

Misgurnus  anguiUicau- 

datus  Cantor 

Branchiae 

Myxoholus  ^. 
CUoromyxum  fujitai 

Nippon 

« 

GaU-bladder 

« 

misgumi 

« 

« 

Myxidium  kagayamai 

« 

« 

Myxoholus  misgumi 

c< 

Molva  vulgaris  Flem 

Bone 

aeglefini 
Leptotheca  informis 

Austria 

GaU-bladder 

Norway 

" 

Sphaeromyxa  hellandi 

« 

MottUa  maculata  Risso 

« 

baJbianii 

France 

M.  tricirrata  BL 

« 

Ceratomyxa  arcuata 
Leptotheca  dongata 

<< 

<i 

",  Monaco 

« 

Sphaeromyxa  halbianii 

France 

« 

sabrasesi 

Monaco 

Mugil  auratus  Risso 

Intestine,  stom- 

ach, spleen, 

pyloric  coeciun 

Myxoholus  miiUeri 

Italy 

Kidney 

exiguus 

" ,  France 

M.  capita  Cuv „ 

« 

exiguus 

<( 

M.  cephalus  L 

GaU-bladder 

Myxidium  incurvatum 

U.  S.  A. 

M.  cheh  Cuv„ 

Stomach,spleen, 

Italy, 

kidney,  etc. 

Myxoholus  exiguus 

France 

2711 


STUDIES  ON  MYXOSPORIDIA—KUDO 


33 


Host 

Organ  Infected 

M3Ttosporidian 

Locality 

M.sp 

Kidney 

Sphaerospora  rostrata 

Italy,  France 
Monaco 

•    i3p 

Muraena  sp 

Gall-bladder 

Ceratomyxa  sp. 

Mustelus  cams  Mitch. 

{M.  vulgaris) 

Gall-bladder 

Ceratomyxa  sphaerulosa 
Myxidium  incurvalum 
Ckloromyxutn  quadratum 

France 

Nerophis  aequoreus  L 

Gall-bladder 

France 

Muscle 

« 

N.  annulatus 

Gall-bladder 

Myxidium  incurvalum 

« 

« 

Sphaeromyxa  sabrazesi 

Monaco 

N.  lumbricifortnis 

« 

Myxidium  incurvalum 
Gen.  et  spec,  incert. 

France 

Notropis  megalops  Raf 

Subcut.  tissue 

U.  S.  A. 

N.  gilberti  J.  et  M 

Muscle 

Myxobolus  orbicttlatus 
aurealus 

U.S.A. 

N,  blennius 

« 
Fins 

« 

N.  anogenus 

C( 

« 

Henneguya  brachyura 

(( 

Oncorkynchus  keta 

Under  the  skin 

Henneguya  salminicola 
« 

Kamtschatk  a 

0,  kisulch 

Connective  tiss. 

« 

of  body  muscle 

« 

Alaska 

0.  nerka 

Gall-bladder 

Chloromyxum  wardi 

Ceratomyxa  arcuata 
Sphaerospora  polymorpha 
Ceratomyxa  arcuata 

« 

Ophidium  vasalli 

Gall-bladder 

Monaco 

Opsanus  tau 

Urin.  bladder 

U.  S.  A. 

Pagellus  centrodontus  Del.... 

Gall-bladder 

France, 

Italy 

Paralichthys  albiguUus 

J.  et  G 

Urin.  bladder 

navicularia 

U.  S.  A. 

^  ^      

« 

spinosa 

« 

« 

Leptotheca  glomerosa 

4( 

« 

Sinuolinea  brachiophora 

(( 

K 

capsularis 

« 

(1 

opacita 

(1 

P.  dentatus 

Gall-bladder 

Ceratomyxa  drepanopsettae 
navicularia 

« 

Urin.  bladder 

(1 

<< 

Leptotheca  lobosa 

C< 

et 

Sinuolinea  capsularis 

« 

Parasilurus  asotus  L 

Intestinal  wall 

Myxobolus  miyairii 
Ceratomyxa  monospora 
Myxobolus  sp. 
Henneguya  wisconsinensis 

Nippon 
U.  S.  A. 

Peprilus  alepidotus 

Gall-bladder 

Perca  flavescens 

Spleen 
Urin.  bladder 

(i 

« 

P.  fluviatilis 

Branchiae 

■    texta 

Italy, 
Germany 

« 

Henneguya  minuta 

« 

« 

Myxobolus  sp. 

Germany 

"  ,  operculum 

permagnus 

« 

(( 

Myxosoma  dujardini 

Switzerland 

« 

Henneguya  psorospermica 

« 

Phoxinus  (Clinostomus) 

Under  scales  on 

fundidoides  Girard 

ext.  surf. 

Myxobolus  transovalis 

U.  S.  A. 

P.laems 

Branchiae 

muJleri 

France 

34 


ILUNOIS  BIOLOGICAL  MONOGRAPHS 


[272 


Host 

Organ  Infected 

Myxosporidian 

Locality 

P.  laevis 

Kidney,  ovary 
Urin.  bladder 
Urin.  bladder 

Gall-bladder 

Membrane  lin- 
ing branchial 
cavity 

Gall-bladder 
? 

Branchiae 

Gall-bladder 
<( 

Otic-capsule 
Muscle 

i( 

GaU-bladder 

Subcutaneous 
muscL  tissue 

Gall-bladder 

« 

« 
« 

Gall-duct 
Gall-bladder 

« 

« 

« 

Branchiae 
Cartilage,  pe- 
richondrium 

Branchiae 
« 

« 

GaU-bladder 

Muscle,  spleen 

Air-bladder 
« 

Gall-bladder 

Myxidium  histophUum 
Sphaerospora  degans 
ZschokkeUa  hildae 

Leptotheca  polymorpha 

Henneguya  linearis 

Myxobolus  notaius 

inaequalis 

Henneguya  linearis 

?  Ceraiomyxa  drepanopsettae 
« 

Sphaerospora  platessae 
Chloromyxum  dupeidae 
dupeidae 
dupeidae 

Ceraiomyxa  acadiensis 
Myxidium  sp. 

Myxobolus  pleuroneciidae 

Chloromyxum  leydigi 
Leptotheca  scissura 
Myxidium  giganteum 
Chloromyxum  leydigi 
Myxidium  sp. 
Chloromyxum  sp. 

Chloromyxum  leydigi 
Chloromyxum  leydigi 

Chloromyxum  leydigi 

Ceratomyxa  sp.  (?) 
Myxobolus  cydoides 

Lentospora  cerebralis 

Myxobolus  cydoides 
scardinii 
Myxosoma  dujardini 

Myxidium  macrocapsulare 

Myxobolus  dispar 

permagnus 
physophilus 

Ceratomyxa  sp.  (?) 

France 

Phycis  blennioides  Br 

P.  mediterraneus  (P. 
i>hvcis  L.) 

Germany 
Norway 

France 

Pimelodus  sebae  Cuv.  et 
Val 

Pimepkales  notatus  Raf 

Piramutana  blochi  C.  et  V. 

Platystoma  fasciatum  L 

Pleuronectes  flesus  L 

S.  America 
Canada 

S.  America 

« 

Norway 
« 

P.  platessa  L 

Pomolobus  aestivalis 

England 
U.  S.  A. 

P.  mediocris  Mitch 

II 

P.  pseudoharengus  Yoxmg.. 
Pseudopleuronedes  amer- 
icanus 

II 

Canada 

Pteroplatea  madura 
Le  Sueur 

(1 

U.S.A. 
<i 

Raja  asterias 

II 

Italy 
France 

R.  bails  L 

R.  radiaia 

Germany  ? 
Murman 

R,  davata  L 

Coast 
France 

R.  undulaia  Lac 

II 

Rkina  squatina  L 

France, 

RMnobaihus  sp.  (?)  Awer.... 
Rhodeus  amarus  BL 

Germany 
Africa 
Germany 

Salmo  foniinalis  Mitch 

Scardinius  erythrophihal- 
mus 

II 
II 

Scatophagus  argus 

Russia 

France, 

Germany 
<i 

France 
Germany 
Russia 
Africa 

2731 


STUDIES  ON  MYXOSPORIDJA—KUDO 


35 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Scoliodon  terrae-novae 

Gall-bladder 

Ceratomyxa  abbreviata 

U.  S.  A. 

(( 

attenuata 

(( 

« 

sphairophora 

(( 

« 

taenia 

(( 

« 

Chloromyxutn  leydigi 

« 

Sconiber  scotnbrus  L 

« 

Leptotheca  parva 

France, 

Norway 

Kidney 

renicola 

France 

? 

Gen.  et  sp.  inc. 

Germany? 

Scorpaena  porcus  L 

Gall-bladder 

Ceratomyxa  arcuata 

France 

S.  scrofd  L 

« 

Ceratomyxa  arcuata 

« 

« 

Myxidium  incurvatum 

« 

S.  sp 

« 

Leptotheca  agilis 

",  Germany 
Monaco 

Scylliwn  canicula 

(( 

Ceratomyxa  sphaerulosa 

<( 

Chloromyxum  leydigi 

" ,  France 
Germany 

S.  dsterids 

« 

a 

Italy 

Norway 

Fastem 

Sebastes  dactylopterus 

(( 

Leptotheca  macrospora 
Leptotheca  sp. 

5.  norvegicus 

« 

Finmark 

5.  viviparus  H.  Kr 

« 

Leptotheca  macrospora 
Spkaeromyxa  sabrazesi 

Norway 
Monaco 

Siphonostoma  rondeletii 

« 

Siphostoma  floridae 

« 

balbianii 

U.  S.  A. 

Urin.  bladder 

Sinuolinea  arborescens 

« 

S.  louisianae 

Gall-bladder 

Spkaeromyxa  balbianii 
Myxidium  gadi 
Ceratomyxa  (?)  spari 
navicularia 

« 

Solea  vulgaris 

« 

France 

Spcrus  berda 

i( 

Africa 

Sphaeroides  tnaculatus 

Urin.  bladder 

U.  S.  A. 

(( 

Sinuolinea  capsularis 

(( 

<( 

Zschokkella  globulosa 

« 

Spinax  spinax  L 

GaU-bladder 

Chloromyxum  leydigi 

Italy, 
France 

Squalis  agassizii  Heck 

Branchiae 

Myxobolus  miilleri 

France 

S.  a.  Savigny  Bona- 

parte  

« 

Myxobolus  miilleri 
Chloromyxum  clupeidae 

(( 

Stenotomus  ckrysops  L 

Muscle 

U.  S.  A. 

Urin.  bladder 

Gen.  et  sp.  incert. 

Canada 

Syngnathus  acus  L 

Gall-bladder 

Myxidium  incurvatum 
Spkaeromyxa  sabrazesi 

France 

« 

Italy, 

Monaco 

Muscle 

Chloromyxum  quadratum 

France 

S.typMe 

GaU-bladder 

Myxidium  incurvatum 

France 

Synodontis  schaU  Bl.  Schn.. 

Integum.  of 

cephalic  reg. 

Henneguya  strongylura 

Egypt 

? 

Myxobolus  inaequalis 

S.  America 

Synodus  faetans 

Gall-bladder 

Ceratomyxa  agglomerata 
amorpha 

U.  S.  A. 

(( 

« 

Tautogolabrus  adspersus 

Walb 

Muscle 

Chloromyxum  clupeidae 

(( 

36 


JLUNOJS  BIOLOGICAL  MONOGRAPHS 


(274 


Host 

Organ  Infected 

Myxosporidian 

Locality 

ThymaUus  thytnallus  L 

Gall-bladder 

Chloromyxum  thymaUi 

Austria 

(( 

Myxobolus  sp. 

« 

Neurilemma  (?) 

pfeifferi 

Germany  ? 

Tkysanophris  japonicus 

Gall-bladder 

Sphaeromyxa  exneri 

Africa 

Tinea  tinea  L.  {T.  vulgaris). 

Branchiae 
Air-bladder, 

Myxobolus  piriformis 

France, 
Germany 

kidney,  etc. 

eUipsoidcs 

(1 

GaU-bladder 

Chloromyxum  cristatum 

France 

<( 

Myxidium  pfeiferi 

Germany 

Kidney 

Myxobolus  cy  print 

Germany, 
Hungary 

Torpedo  narce  Risso 

Gall-bladder 

Chloromyxum  leydigi 
Chloromyxum  leydigi 
Chloromyxum  leydigi 
"Ceratomyxa  reticularis 
Myxidium  incurvatum 

France 

T.  oceUata 

(( 

Germany 

T.  torpedo  L 

(( 

Trachinus  draco  IL 

« 

« 

(( 

« 

Trackurus  trachurus  L 

Muscle 

Chlordmyxum  quadrcUum 

France, 
Germany 

Trutta  fario  L 

Gall-bladder, 

Gall-duct 

Chloromyxum  truttae 

France 

Nervous  syst. 

Myxobolus  neurobius 

Germany 

Subcutaneous 

conn.  tiss.  at 

"• 

base  of  fin 

Henneguya  niisslini 

« 

T.  iridea  Gibb 

Cartilage,  peri- 
chondrium 

Lentospora  eerebralis 

« 

T.  solar  L 

« 
Gall-bladder 

'                               cc 

Myxidium  oviforme 

« 

Norway 

Trygon  pastinaca  L 

Gall-bladder 

Chloromyxum  leydigi 
Leptotheca  agilis 

France 

"  ,  Italy 

Tylosurus  inarianus 

Urin.  bladder 

Chloromyxum  granulosum 
Ceratomyxa  tylosuri 

U.  S.  A. 

T.  scMsmatorhynckus 

Gall-bladder 

Africa 

Urophycis  ckuss 

« 

acadiensis 

Canada 

Zoarees  angularis 

« 

acadiensis 

« 

Amohibia              -• 

Kidney 
Gall-bladder 

Wardia  ohlmacheri 

Bufo  lentiginosus 

U.  S.  A. 

B.  marinus  L 

Sphaeromyxa  immersa 
Myxobolus  hylae 
Sphaeromyxa  immersa 

Brazil 

Hyla  aurea 

Testis,  ovary 
GaU-bladder 

Australia 

Leptodactylus  oceUaius 

Brazil 

Molge  cristata  Laur. 

(Tritonc.) ; 

« 

Chloromyxum  caudatum 

France 

Proteus  anguineus  L 

Kidney 
« 

Chloromyxum  protei 
?Wardia  ohlmacheri 
?Wardia  ohlmacheri 

Austria 

Rana  esctdenta 

France? 

R.  temporaria  (R.fusca) 

* 

Reptilia 

Emys  orbicularis  L. 

{Cistudo  europaea) 

Kidney 

Myxidium  danilewskyi 

Russia, 

France 

2751 


STUDIES  ON  MYXOSPORJDIA—KUDO 


37 


Host 

Organ  Infected 

Myxosporidian 

Locality 

Lacerta  sp 

Ovarian  egg 
Kidney 

Gen.  et  spec,  incert. 

Myxidiutn  mackiei 

americanum 

Italy 
India 

Trionyx  (Amyda)  ganzeti- 
cus 

T.  spinifera 

U.  S.  A. 

C.  DISTRIBUTION  OF  MYXOSPORIDIA  IN  THE  ORGANS  OF  THE  HOST 

Altho  some  species  are  found  in  various  organs  of  the  host  animal,  the 
majority  has  one  or  two  particular  seats  of  infection.  Among  the  various 
organs  which  become  infected,  the  gall-bladder  is  most  frequently  infected. 
The  kidney,  branchia  and  urinary  bladder  have  less  chances  of  being 
parasitized.  As  to  the  infection  of  the  reproductive  organs  of  the  host, 
little  is  known.  The  male  reproductive  organ  becomes  rarely  infected, 
being  reported  only  twice.  The  female  reproductive  organ,  however,  is 
more  frequently  infected.  The  infection  of  the  next  generation  of  the  host 
animal  thru  the  infected  ovum  which  is  known  to  occur  in  some  Micro- 
sporidian  parasites,  has  not  been  reported  in  Myxosporidia  as  yet. 

LIST  IV.    ORGANS  OF  HOST  INFECTED  BY  MYXOSPORIDIA 


I.  Pisces 

1)  Integument. — Sphaerospora  sp.  Southwell  et  Prashad  (under  the  scales) 

Myxobolus  seni  Southwell  et  Prashad  (fin) 

Myxobolus  transovdis  Gurley  (under  the  scales) 

Myxobolus  unicapsulatus  Gurley  (head) 

Myxobolus  cycloides  Gurley  (opercle) 

Myxobolus  inaequalis  Gurley  (head) 

Myxobolus  sp.  Gurley  (opercle,  head,  fin) 

Myxobolus  squamae  Keysselitz  (inner  surface  of  the  scales) 

Myxobolus  volgensis  Reuss  (fin) 

Myxobolus  pertnagnus  Wegener  (operculum) 

Myxobolus  aureatus  Ward  (fin) 

Henneguya  brachyura  Ward  (fin-ray) 

Henneguya  linearis  (Gurley)  Labb^  (membrane  lining  branchial  cavity) 

Henneguya  gurleyi  Kudo  (base  of  spines  of  dorsal  fin) 

Henneguya  strongylura  (Gurley)  Labb6  (cephalic  region) 

Lentospora  dermatobia  Ishii 

2)  Connective  tissue. — Myxidiutn  anguillae  Ishii  (subcutaneous) 

Myxobolus  fuhrmanni  Auerbach  (under  the  oral  mucous  membrane) 

Myxobolus  oviformis  Th61ohan  (subcutaneous) 

Myxobolus  lintoni  Gurley  (subcutaneous) 

Myxobolus  oblongus  Gurley  (chiefly  of  the  head) 

Myxobolus  gigas  Auerbach  (of  operculum,  sides  and  fins) 

Myxobolus  capsulatus  Davis  (visceral) 

Henneguya  kolesnikovi  (Gurley)  Labb6  (interstitial) 

Henneguya  niisslini  Schuberg  et  Schroder  (at  the  base  of  dorsal  fin) 

Henneguya  miyairii  Kudo  (of  the  head) 

Gen.  et  sp.  incert.  Linton  (subcutaneous) 


38  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [276 

3)  Muscle. — Leptolheca  perlata  (Gurley)  Labb£ 

Chloromyxum  quadratum  Th^lohan 

Chloromyxum  fttnduli  Hahn 

Chloromyxum  clupeidae  Hahn 

Lentospora  multiplicata  Reuss 

Myxoboltis  notatus  Mavor  (connective  tissue  of  voluntary  muscle) 

Myxoholus  pfeifferi  Tli6lohan 

Myxobolus  sandrae  Reuss 

Myxoholus  musculi  Keysselitz 

Myxobolus  funduli  Kudo 

Myxobolus  pleuronectidae  Hahn 

Myxobolus  sp.  Southwell  (subcutaneous  intermuscular  tissue) 

Myxobolus  nodularis  Southwell  et  Prashad 

Myxobolus  orbiculatus  Kudo 

Henneguya  creplini  (Gurley)  Labb€ 

Henneguya  monura  (Gurley)  Labbfi 

Henneguya  zschokkei  (Gurley)  Doflein 

Henneguya  salminicola  Ward 

Gen.  et  spec,  incert.  Leydig 

4)  Eye. — Sphaerospora  platessae  Woodcock  (optic  capsule) 

Myxobolus  oculi4eucisci  Trojan  (vitreous  body) 

Myxobolus  ellipsoides  Thflohan 

Myxobolus  miiUeri  Biitschli 

Myxobolus  aeglefini  Auerbach 

Myxobolus  volgensis  Reuss 

Myxobolus  magnus  Awerinzew 

Henneguya  sckizura  (Gurley)  Labb€  (intercellular  tissue  of  eye  muscle) 

5)  Branchiae. — Sphaerospora  carassii  Kudo 

Myxosoma  dujardini  Th61ohan 

Myxosoma  (?)  lobatum  Nemeczek 

Myxosoma  funduli  Kudo 

Lentospora  acuta  (Fujita) 

Myxobolus  piriformis  Th^lohan 

Myxobolus  toyamai  Kudo 

Myxobolus  rohitae  Southwell  et  Prashad 

Myxobolus  dispar  Thfilohan 

Myxobolus  ellipsoides  Th^lohan 

Myxobolus  exiguus  Thdlohan 

Myxobolus  oviformis  Thfilohan 

Myxobolus  miilleri  Biitschli 

Myxobolus  globosuS  Gurley 

Myxobolus  cycloides  Gurley  (also  pseudobranchiae) 

Myxobolus  sphaeralis  Gurley 

Myxobolus  anurus  Cohn 

Myxobolus  sp.  Gurley 

Myxobolus  gigas  Auerbach 

Myxoboltis  volgensis  Reuss 

Myxobolus  scardinii  Reuss 

Myxobolus  macrocapsidaris  Reuss 

Myxobolus  bramae  Reuss 

Myxobolus  cyprinicola  Reuss 


277]  STUDIES  ON  MYXOSPORIDIA— KUDO  39 

Myxobolus  balleri  Reuss 
Myxobolus  sp.  Miyairi 
Myxobolus  sp.  Wegener  • 

Myxobolus  perntagnus  Wegener 
Myxobolus  rotundus  Nemeczek 
Myxobolus  minutus  Nemeczek 
Myxobolus  funduli  Kudo 
Myxobolus  koi  Kudo 
Myxobolus  discrepans  Kudo 
Henneguya  psorospermica  Th6lohan 
Henneguya  texla  (Cohn) 
Henneguya  minuta  (Cohn) 
Henneguya  lobosa  (Cohn) 
Henneguya  creplini  (Gurley)  Labb6 
Henneguya  linearis  (Gurley)  Labb6 
Henneguya  acerinae  Schroder 
Henneguya  gigantea  Nemeczek 
Gen.  et  spec,  incert.  Heckel  et  Kner 

6)  Heart. — Myxobolus  cordis  Keysselitz  (muscle  of  ventricle  and  bulbus  arteriosus) 

Gen.  et  spec,  incert.  Leydig  (auriculo-ventricular  valve) 

7)  Air  bladder. — Myxobolus  ellipsoides  Th^lohan  (conn,  tiss.) 

Myxobolus  miilleri  Butschli  (conn,  tiss.) 
Myxobolus  physophilus  Reuss  (sxirface) 
Myxobolus  perntagnus  Wegener 

8)  Body-cavity  (cyst). — Myxobolus  sp.  Gurley 

Myxobolus  carassii  Klokacewa 
Gen.  et  spec,  incert.  Leydig 

9)  Nervous  tissue. — Myxobolus  neurobius  Schuberg  et  Schroder 

Lentospora  encephalina  Mulsow  (blood  vessel  of  brain) 

10)  Bone,  cartilage,  perichondrium. — Lentospora  cerebralis  (Hofer)  Plehn 

Myxobolus  aeglegini  Auerbach 
Henneguya  brachyura  Ward  (of  fin) 

11)  Stomach,  pyloric  cecum. — Myxobolus  exiguus  Thfilohan 

Myxobolus  mesentericus  Kudo 
Henneguya  tenuis  Vaney  et  Contc 

12)  Liver. — Myxobolus  ellipsoides  Th61ohan 

Myxobolus  oviformis  Th^lohan 
Myxobolus  cyprini  Doflein 
Myxobolus  cordis  Keysselitz 
Myxobolus  musculi  Keysselitz 
Myxobolus  carassii  Ellokacewa 
Myxobolus  mesentericus  Kudo 

13)  Gall-bladder. — Leptoiheca  agilis  Th^lohan 

Leptotheca  elongata  Thfilohan 
Leptotheca  polymorpha  (Th61ohan)  Labb6 
Leptotheca  parva  Th61ohan 
Leptotheca  hepseti  Th61ohan 
Leptotheca  sp.  Awerinzew 
Leptotheca  macrospora  Auerbach 
Leptotheca  informis  Auerbach 
Leptotheca  longipes  Auerbach 


40  ILUNOIS  BIOLOGICAL  MONOGRAPHS  1278 

Leptotheca  fusifonnis  Davis 
LepMheca  scissura  Davis 
Caratomyxa  arcuaia  Thflohan 
Ceratomyxa  spkaerulosa  Th^lohan 
Ceratomyxa  paUida  Th61ohan 
Ceratomyxa  globurijera  Thelohan 
Ceratomyxa  appendiadata  Th61ohan 
Ceratomyxa  truncata  Thelohan 
Ceratomyxa  reticularis  Th6lohan 
Ceratomyxa  inaequalis  Doflein 
Ceratomyxa  linospora  Doflein 
Ceratomyxa  ramosa  Awerinzew 
Ceratomyxa  drepanopsettae  Awerinzew 
Ceratomyxa  tylosuri  Awerinzew 
Ceratomyxa  (?)  spari  Awerinzew 
Ceratomyxa  sp.  (?).  Awerinzew 
Ceratomyxa  sp  (?).  Awerinzew 
Ceratomyxa  acadiensis  Mavor 
Ceratomyxa  sp.  Georgdvitch 
Ceratomyxa  coris  Georgfivitch 
Ceratomyxa  herouardi  Georg6vitch 
Ceratomyxa  mesospora  Davis 
Ceratomyxa  sphairophora  Davis 
Ceratomyxa  taenia  Davis 
Ceratomyxa  attenuata  Davis 
Ceratomyxa  recurvata  Davis 
Ceratomyxa  lunaia  Davis 
Ceratomyxa  abbreviata  Davis 
Ceratomyxa  flagellif era  Davis 
Ceratomyxa  agglomerata  Davis 
Ceratomyxa  amorpha  Davis 
Ceratomyxa  monospora  Davis 
Ceratomyxa  streptospora  Da\'is 
Ceratomyxa  aggregata  Davis 
Ceratomyxa  undidata  Davis 
CUoromyxum  leydigi  Mingazzini 
CUoromyxum  fluviatile  Th61ohan 
CUoromyxum  truttae  L6ger  (also  gall-duct) 
CUoromyxum  cristatum  Ldger 
CUoromyxum  dubium  Auerbach 
CUoromyxum  sp.  Awerinzew 
CUoromyxum  thymaUi  Lebzelter 
CUoromyxum  koi  Fujita 
CUoromyxum  magnum  Awerinzew 
CUoromyxum  misgumi  Kudo 
CUoromyxum  fujiiai  Kudo 
CUoromyxum  trijugum  Kudo 
CUoromyxum  catostomi  Kudo 
CUoromyxum  wardi  Kudo 
Sphaerospora  masovica  Cohn 
Myxidium  incurvatum  Thflohan 
Myxidium  sphaericum  Thdohan 


2791  STUDIES  ON  MYXOSPORIDJA—KUDO  41 

Myxidiutn  sp.  Guriey  (only  in  gall-duct) 

Myxidium  giganteum  Doflein 

Myxidium  pfei£eri  Auerbach 

Myxidiutn  inflatum  Auerbach 

Myxidium  bergense  Auerbach 

Myxidiutn  procerutn  Auerbach 

Myxidiutn  tnacrocapsulare  Auerbach 

Myxidiutn  sp.  Awerinzew 

Myxidiutn  ovifortne  Parisi 

Myxidium  sp.  Mavor 

Myxidium  kagayamai  Kudo 

Myxidium  gadi  Georg6vitch 

Myxidium  glutinosum  Davis 

Myxidium  phyllium  Davis 

Myxidium  striatum  Cunha  et  Fonseca 

Myxoholus  misgurtii  Kudo  (few  spores  only) 

Myxobolus  sp.  Lebzelter  (spores  only) 

Sphaeromyxa  balbiatiii  Th61ohan 

Sphaeromyxa  incurvata  Doflein 

Sphaeromyxa  sabrazesi  Laveran  et  Mesnil 

Sphaeromyxa  hellandi  Auerbach 

Sphaeromyxa  exneri  Awerinzew 

Sphaeromyxa  gasterostei  Georgdvitch 

Zschokkella  twva  Klokacewa 

Zschokkella  acheilognathi  Kudo  (also  in  gall-duct) 

Gen.  incert.  congri  Perugia 

Gen.  incer.  merlucii  Perugia 

Gen.  et  spec,  incert.  Mavor 

14)  Spleen. — Myxobolus  piriformis  Th^lohan 

Myxobolus  sp.  Kudo 
Myxobolus  ellipsoides  Th61ohan 
Myxobolus  exiguus  Thdohan 
Myxobolus  omformis  Th^lohan 
Myxobolus  pfeifferi  Th^lohan 
Myxobolus  cyprini  Doflein 
Myxobolus  cordis  Keysselitz 
Myxobolus  musculi  Keysselitz 
Myxobolus  mesenlericus  Kudo 

15)  Intestine. — Myxobolus  exiguus  Thelohan 

Myxobolus  miilleri  Biitschli 
Myxobolus  pfeifferi  Th61ohan 
Myxobolus  miyairii  Kudo 
Myxobolus  carassii  Klokacewa 
Myxobolus  mesenlericus  Kudo 
Hentieguya  peri-intestinalis  C6pSde 
Henneguya  tenuis  Vaney  et  Conte 

16)  Ovary.^Wardia  ovinocua  Kudo 

Sphaerospora  elegans  Th61ohan 
Myxidium  histophilum  Th61ohan 
Myxobolus  pfeifferi  Thelohan 
Myxobolus  miilleri  Biitschli 
Myxobolus  musculi  Keysselitz 


42  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [280 

Eenneguya  oviperda  (Cohn) 
Henneguya  media  Thflohan 
Eenneguya  brevis  TMlohan 

17)  Kidney. — a)  Urinary  tubules. — Leptotheca  renicola  Thdlohan 

Mitraspora  caudata  (Parisi)  Kudo 
Mitraspora  cyprini  Fujita  (also  in  ureter) 
Mitraspora  elongata  Kudo 
Sphaerospora  elegans  Thfilohan 
Sphaerospora  divergens  Th^ohan 
Eenneguya  media  Th61ohan 
Eenneguya  brevis  Th61ohan 
Eenneguya  gasterostei  Parisi 
Eoferellus  cyprini  Doflein 

b)  Tissue. — Mitraspora  elongala  Kudo 

CUoromyxum  quadratum  Thelohan 

Sphaerospora  rostrata  Thelohan  (^lalpighian  bodies) 

Myxidium  kistophilum  Th61ohan 

Lentospora  asymmetrica  Parisi 

Myxobolus  pfeifferi  Tli61ohan 

Myxobolus  cyprini  Doflein 

Eenneguya  neapolitana  Parisi  (conn.  tiss.  of  ren.  tubules) 

Eoferellus  cyprini  Doflein 

c)  Seat,  unstated. — CUoromyxum  mucronaium  Gurley 

Sphaerospora  angtdata  Fujita 
Myxidium  barbatulae  CdpMe 
Myxidium  giardi  C^pede 
Myxobolus  piriformis  Thelohan 
Myxobolus  ellipsoides  Thdlohan 
Myxobolus  exiguus  Thelohan 
Myxobolus  oviformis  Thelohan 
Myxobolus  mOUeri  Btitschli 
Myxobolus  obesus  Gurley 
Myxobolus  cycloides  Gurley 
Myxobolus  cordis  Keysselitz 
Myxobolus  musculi  Keysselitz 

18)  Urinary  bladder. — Leptotheca  lobosa  Davis 

Leptotheca  glomerosa  Davis 
Ceratomyxa  naricularia  Davis 
Ceratomyxa  spinosa  Davis 
CUoromyxum  mucronaium  Gurley 
CUoromyxum  granulosum  Davis 
Sphaerospora  elegans  Thelohan 
Sphaerospora  divergens  Th61ohan 
Sphaerospora  polymorpha  Davis 
Sphaerospora  sp.  Davis 
Sinuolinea  dimorpha  Davis  (also  in  ureter) 
Sinuolinea  capsularis  Davis 
Sinuolinea  arborescens  Davis 
Sinuolinea  opacHa  Davis 
Sinuolinea  brachiophora  Davis 
Myxidium  libber kiihni  Biitschli 
Zschokkella  hildae  Auerbach 
Zschokkella  globulosa  Davis 


281] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


43 


Myxoproteus  atnbiguus  (Th^lohan)  Doflein 

Myxoproteus  cordiformis  Davis 

Myxoproteus  cornutus  Davis 

Henneguya  legeri  C6pSde 

Henneguya  wisconsinensis  Mavor  et  Stxasser 

Henneguya  mictospora  Kudo 

Gen.  et  spec,  incert.  Mavor 

19)  Testis. — Myxobolus  pfeifferi  Th61ohan  (only  spores) 

20)  Mesentery. — Myxobolus  mesentericus  Kudo 

21)  Seat  unknown. — Henneguya  sp.  Gurley  (integument?) 

Gen.  et.  spec,  incert.  Borne 

II  Amphibia 

1)  Gall-bladder. — CMoromyxum  caudatutn  Th^Iohan 

Sphaeromyxa  immersa  (Lutz)  Thfilohan 

2)  Urinary  tubules  of  kidney. — Wardia  oUmacheri  (Gurley)  Kudo 

CMoromyxum  protei  Joseph 


TABLE  I 

4-1 

a 

a 

Si 

1 

o 

3 

< 

i 

'•3 

i 

1 

M 

O 
CO 

t3 

O 
11 

33 

3, 

C/5 

=3 

1 

5 

.2 

O 

1 

1(a)* 

1 

rtT3 
t33 

2 
2 
3 

b 

Si 

a 

o 
w  9 

Leptotheca 

Ceratomyxa 

Myxoproteus.... 

Wardia 

1 

1(a) 

3(a) 

Kb)* 

1(a) 

Kb) 

1(c)* 

2(a) 

Kb) 

1(c) 

2(a) 

Kb) 

2(c) 

3(a) 

Mitraspora 

3 

14 

1 

Chlorom3rxum .. 

1 

2 

4 
5 

1 

1 

1 

1 

1 

Sphaerospora.... 

Sinuolinea 

Myxidium 

1 

18 

7 
2 

1 

Sphaeromyxa.... 

Zschokkella 

2 

Myxosoma 

3 
1 

28 

8 

Lentospora 

1 

10 
4 

6 

3 

1 
9 

4 

6 
1 

1 

4 

1 
1 

1 
1 

1 

Kb) 
2(b) 
9(c) 
3(a) 
Kb) 
1(a) 
Kb) 

1 

1 

Myxobolus 

Henneguya 

2 

1 

7 

2 

10 

6 
2 

2 

4 
3 

3 

1 

....  1 
1 

Hoferellus 

Total 

16 

10 

18 

8 

41 

1 

4 

9 

^ 

^ 

7 

88 

10 

s 

2 

10 

39 

3 

24 

1 

7 

'  For  (a),  (b)  and  (c),  see  page  42. 


44 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[282 


3)  Testis,  oviduct. — Myxobcius  hylat  Johnston  ct  Bancroft 

m  RZPTILIA 

1)  Kidney  (ren.  tub.). — Myxidium  danilewskyi  Laveran 

Myxidium  mackiti  Bosanquet 
Myxidium  atnericanum  Kudo 

2)  Ovary. — Gen.  et  ^)ec.  incert.  Mingazzini 

rV  Insecta 
1)  Abdominal  cavity. — Chhromyxum  diploxys  Guriey 

V  Annelida 
Myxoboius  ^.  Guriey 

The  data  in  this  section  are  summarized  on  the  preceding  page  (Table  I). 


D.    THE  EFFECT  OF  ENVIRONMENT  ON  THE  ORGANAL  DISTRIBUTION 
OF  MYXOSPORmL\  IN  HOSTS 

Myxosporidia  are  almost  equally  distributed  among  marine  and  fresh- 
water fishes  in  regard  to  the  number  of  species.  This  is  shown  in  the  follow- 
ing table. 

T.\BLE  II 


Number  of  species 

Number  of  species 

Other  hosts 

Genus 

found  in  marme 

found  in  fresh- 

fi»h 

water  fish 

RepL 

Amph. 

In.sect 

Annelida 

Leptotheca  (15)*.„. 

15 

Ceratomyxa  (35).... 

35 

Myxoproteus  (3).... 

3 

Wardia  (2) 

1 

1 

Mitraspora  (3) 

3 

.... 

.... 

Ghloromjnum  (22) 

7 

12 

2 

.... 

Sphaerospora  (10). 

5 

5 

.... 

Sinuolinea  (5) 

5 

.... 

Myxidium  (26) 

17 
(2  common) 

8 
(2  common) 

3 

.... 

Sphaeromyxa  (7).... 

6 

.... 

.... 

1 

.... 

ZschokkeUa  (4) 

2 

2 

.... 

.... 

Mjniosoma  (3) 

1 

2 

.... 

.... 

Lentospora  (6) 

2 

(2  common) 

6 

(2  common) 

.... 

Mjrxobolus  (63) 

5 

56 

1 

1 

Henneguya  (32) 

1 

31 

HofereUus  (1) 

.... 

1 

.... 

.... 

Gen.  et  spec 

incert  (12) 

4 

7 

1 

.... 

.... 

Total  237+12 

104+4 

134+7 

4 

5 

1 

1 

*  The  number  in  parenthesis  denotes  the  number  of  species  in  the  corresponding  genus. 


2831 


STUDIES  ON  MYXOSPORJDJA—KUDO 


45 


These  genera  have  certain  relations  to  the  organal  distribution  in  the 
body  of  the  host,  which  are  shown  in  List  IV  (page  37)  and  Table  I  (on  page 
43)  and  which  can  be  put  together  as  follows : 


TABLE  III 


Genus 

Number  of  species 
found  in  body- 
cavity 

Number  of  species 
found  in  tissue 

Number  of 

species  found 

in  both 

Seat 
unknown 

Leptotheca  (15) 

14 

35 
3 
1 
2 

18 

'  4 

5 

22 
6 
4 

1 
2 
4 

4 

1 

1 

4 
4 

4 

3 

5 

59 

28 

5 

1 

1 

1 

Ceratomyxa  (35) 

Myxoproteus  (3) 

Wardia  (2) 

Mitraspora  (3) 

Chloromyxum  (22) 

Sphaerospora  (10) 

Sinuolinea  (5) 

Myxidium  (26) 

Sphaeromyxa  (7) 

Zschokkella  (4) 

Myxosoma  (3) 

Lentospora  (6) 

Myxobolus  (63) 

Henneeuva  (32) 

Hoferellus  (1) 

Gen.  et  spec,  inct  (12).... 

3 

Total  237+12 

121+4 

109+5 

3 

4+3 

From  the  facts  shown  in  the  above  tables,  the  following  conclusions  can 
be  drawn. 

1)  The  genera  Leptotheca  (one  species  in  tissue),  Ceratomyxa,  Myxo- 
proteus, Sinuolinea  and  Sphaeromyxa  (one  species  in  Amphibia)  include 
parasites  from  the  body  cavity  of  marine  fish. 

2)  The  majority  of  the  genera  Lentospora,  Myxosoma,  Myxobolus  (one 
species  in  Amphibia),  Henneguya  and  Hoferellus  include  parasites  in  tissues 
of  fresh-water  fish. 

3)  The  genera  Chloromyxum,  Sphaerospora,  Myxidium  and  Zschok- 
kella include  forms  that  infect  the  body-cavity  as  well  as  the  tissue  of 
marine  and  fresh-water  fishes. 

4)  The  genus  Mitraspora  includes  three  species  that  parasitize  the 
fresh-water  fish. 

5)  The  genus  Myxidium  has  three  species  found  in  the  kidney  of 
reptiles. 


46  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [284 

6)  The  genera  Wardia,  Chloromyxum,  Sphaeromyxa  and  Myxobolus 
include  species  parasitic  in  Amphibia. 

7)  The  genus  Chloromyxum  has  one  species  found  in  an  insect. 

8)  The  genus  Myxobolus  has  one  species  found  in  an  annelid,  which 
was  not  normally  recorded. 


285] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


47 


THE  SPORE 

As  will  be  shown  later  (page  52-55),  the  spore  stage  is  still  the  only 
constant  character  by  which  various  forms  of  Myxosporidia  are  identified 
from  each  other.  For  this  reason  it  is  necessary  to  have  a  clear  conception 
of  the  form  and  structure  of  the  spore  and  at  the  same  time  to  define  the 
terms  used  in  the  present  paper,  even  tho  they  have  commonly  been 
used  heretofore. 

Anterior  end 
Foramen  or  polar  capsule 

Shell 
sutural  ridge 

Polar  capsule 
Coiled  polar  filament 


Nuclei  07  sporoplasm 
Sporoplasm 

lODINOPHlLOUS  vacuole 
POSTXKIOX  END 

A  B 

Textfigure 
Diagrammatical  front  [A]  and  side  p]  views  of  a  Mvxoboltjs  spore. 

(For  further  explanation  see  following  pages.) 

The  spore  of  Myxosporidia  is  covered  by  a  shell,  which  is  composed 
of  two  valveSy  usually  symmetrical  in  form  and  size  that  come  in  contact 
in  the  sutural  plane.  The  sutural  line  is  straight  in  most  cases,  tho  some- 
times curved  like  an  S.  It  is  more  or  less  thickened,  forming  the  sutural 
ridge.  The  sutural  ridge  is  to  be  made  out  clearly  in  fresh  as  well  as  in 
stained  preparations  and  furnishes  important  data  in  regard  to  the  classi- 
fication of  the  parasite.  The  thickness  of  the  shell- valve  is  usually  uniform; 
in  some  species  (Myxobolus),  however,  it  may  differ  slightly  in  different 
parts  of  the  shell.  Besides,  in  many  species  of  Myxobolus,  the  shell 
differentiates  a  small  triangular  intercapsular  appendix  on  the  inside  at  the 
anterior  end  directed  posteriad  between  two  polar  capsules. 

The  form  of  the  spore  varies  greatly  owing  to  the  shape  of  the  shell 
together  with  its  variously  developed  appendages;  1)  lateral  appendages 
as  in  Ceratomyxa,  2)  anterior  processes  as  in  Myxoproteus,  3)  posterior 
processes  as  in  Wardia  (fringe-like),  Mitraspora  (filiform),  Hoferellus 
(spinous),  Henneguya  (tail-form),  etc. 


48  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [286 

The  surface  of  the  shell  may  be  smooth  or  exhibit  various  markings. 
More  or  less  conspicuous  ridges  varying  in  form  and  number  in  different 
species,  may  run  parallel  to  the  sutural  line,  may  show  a  network-like 
structure  or  may  exhibit  short  tooth-like  processes  arising  from  the  sutural 
ridge  and  radiating  toward  the  center  of  each  valve.  When  the  ridges  are 
fine,  they  form  delicate  striations,  arranged  usually  parallel  to  the  sutural 
line.  Tho  these  markings  are  usually  easily  seen  in  vivo,  they  are  very 
often  more  readily  studied  in  stained  preparations. 

Inside  of  the  shell  are  present  the  polar  capsules  and  sporoplasm. 
Gurley  (1894:120)  and  Davis  (1917:210)  used  the  term  "capsules"  instead 
of  polar  capsules  because  of  the  facts  that  "the  situation  implied  by  the 
the  latter  (polar  capsule)  is  not  constant"  (Gurley)  and  that  "they  are 
often  not  in  the  position  indicated  by  the  term  polar  capsule"  (Davis). 
The  present  writer,  however,  does  not  agree  with  these  authors  and  retains 
the  commonly  used  term,  polar  capsule,  thruout  the  present  paper  on 
the  basis  of  the  fact  that  these  polar  capsules  are  situated  at  or  near  the 
more  or  less  attenuated  anterior  end  in  the  great  majority  of  species  or  at 
each  end  (in  Myxidiidae)  of  the  spore,  except  in  the  few  cases  as  in  Wardia 
in  which  they  are  situated  in  the  central  portion  and  have  the  foramina  at 
the  anterior  end  of  the  spore. 

The  polar  capsules  may  be  pyriform  or  spherical.  They  are  located 
at  or  near  one  end  (anterior  end)  of  the  spore.  In  Myxidiidae,  one  polar 
capsule  is  situated  at  each  end,  in  which  case  no  distinction  can  be  made 
between  the  anterior  and  posterior  ends.  The  end  or  side  opposite  to  the 
anterior,  is  the  posterior  end  of  the  spore.  The  number  of  polar  capsules 
in  a  spore  varies  according  to  the  dififerent  genera.  There  is  only  one  polar 
capsule  in  the  spore  of  unicapsular  Myxobolus,  four  in  Chloromyxum,  two 
in  all  the  other  genera.  They  may  be  equal  or  unequal  in  form  and  size. 
When  two  polar  capsules  are  located  at  the  anterior  end,  they  may  be 
convergent  or  divergent.  Each  has  a  foramen  to  the  outside  of  the  spore 
thru  the  shell  in  or  near  the  sutural  line,  thru  which  the  polar  filament  is 
extruded.  The  foramen  is  observable  in  the  fresh  condition.  Staining 
will  very  often  show  clearly  the  canals  thru  the  shell.  Each  polar  capsule 
has  an  independent  foramen. 

In  the  polar  capsule  exists  a  coiled  polar  filament,  which  in  most 
cases  can  be  recognized  without  diflficulty  in  the  fresh  condition.  The 
polar  filament  is  as  a  rule  a  more  or  less  extended,  probably  hollow  thread 
connected  with  the  polar  capsule,  which  is  extruded  from  the  spore  thru 
the  foramen  under  the  action  of  the  stimulants  such  as  the  digestive  fluid  of 
the  host  or  certain  chemicals.  In  Sphaeromyxa  it  is  rather  short  and  thick, 
tapering  to  a  point.  The  polar  filament  is  coiled  around  the  longest 
axis  of  the  polar  capsule,  except  in  Sphaeromyxa  in  which  it  is  coiled 
around  an  axis  perpendicular  to  the  longest  axis  of  the  polar  capsule. 


287]  STUDIES  ON  MYXOSPORIDIA— KUDO  49 

The  sporoplasm  occupies  the  extracapsular  cavity  at  the  posterior 
region  of  the  spore.  It  is  of  granular  structure  with  almost  always  two 
nuclei.  Besides,  it  has  an  iodinophilous  vacuole  mostly  round  or  oval  in 
the  spores  of  the  family  Myxobolidae.  It  occurs  thruout  the  spore  stage 
and  is  the  important  character  of  the  said  family.  The  contents  of  the 
vacuole  is  probably  of  glycogenous  nature  and  is  stained  deeply  with 
iodine.    Small  refringent  fat  globules  have  also  been  observed  in  the  spore. 

Davis  (1917:212)  proposed  to  use  capsular  and  postcapsular  sides  in 
place  of  anterior  and  posterior  ends  which  have  most  frequently  been 
used  and  are  also  used  in  the  present  paper.  The  latter  terms  can  be 
employed  as  properly  as  Davis'  terms  except  in  the  case  of  the  Myxidiidae, 
where  both  terms,  strictly  speaking,  are  inapplicable. 

Tho  various  abnormal  spores  are  very  often  encountered  in  several 
species,  the  majority  of  the  spores  are  of  typical  form,  structure  and  size. 
In  Myxosoma  and  Myxobolus,  the  spore  sometimes  develops  a  short 
posterior  process,  which  is  highly  developed  in  the  spore  of  the  genus 
Henneguya. 

Young  spores,  generally  speaking,  are  more  rounded  in  form  than  the 
mature  form,  while  the  mature  spores,  as  a  rule,  are  of  definite  form, 
structure  and  size  characteristic  to  the  species.  It  should,  however,  be 
kept  in  mind  that  there  is  a  certain  amount  of  variation  among  these 
characters. 

As  is  generally  recognized,  one  must  mention  whether  the  spores  were 
measured  in  fresh  condition  or  in  fixed  and  stained  state.  The  fresh  spore 
is  generally  more  or  less  larger  than  the  mounted  one. 


so  LUNOIS  BIOLOGICAL  MONOGRAPHS  [288 


DEFINITION  OF  TERMS  USED  FOR  DESCRIPTIONS 

Anterior  end. — The  end  of  the  spore  where  the  polar  capsules  open;  in  most 

cases  the  polar  capsules  are  situated  at  this  end. 
Anterior  process. — The  spinous  process  of  the  shell  at  the  anterior  end  of 

the  spore  of  the  genus  Myxoproteus. 
Breadth. — The  larger  diameter  of  the  spore  measured  at  right  angles  to  the 

length  or  sutural  diameter;  the  shorter  diameter  thus  measured  being 

the  thickness. 
Capsulogenous  cell. — A  small  island  of  protoplasm  with  a  nucleus,  in  which 

polar  capsule  becomes  differentiated. 
Cyst. — The  vegetative  form  of  more  or  less  conspicuous  size  in  tissues  of  the 

host,  surrounded  usually  by  a  membranous  structure  composed  of  the 

host  issue. 
Disporous. — The  character  of  a  trophozoite  of  forming  only  two  spores. 
Foramen. — Opening  of  the  polar  capsule*  thru  which  the  polar  filament  is 

extruded. 
Front  view. — The  view  in  which  length  and  breadth  of  the  spore  are  laid 

horizontally. 
Gemmules. — A  small  mass  of  trophozoite  separated  from  the  mother  body 

by  plasmotomy.    Used  by  Davis  (1917).    (See  page  105.) 
lodinophilous  vacuole. — The  vacuole  in  the  sporoplasm  of  the  spore  of  the 

family  Myxobolidae,  the  contents  of  which  are  stained  brownish  with 

iodine. 
Lateral  process. — The  lateral  prolongation  of  the  shell-valve  at  right  angles 

to  the  sutural  plane. 
Length. — Antero-posterior  diameter  of  the  spore  in  the  sutural  plane; 

equivalent  to  sutural  diameter.  ♦ 

Longitudinal  striations. — Fine  ridges  or  thickenings  marked  longitudinally 

on  the  shell  of  the  spore. 
Mesoplasm. — An  intermediate  layer  between  ectoplasm  and  endoplasm, 

coined  by  Cohn  in  the  case  of  Myxidium  lieherkiihni  (see  page  107). 
Mictosporous. — The  character  of  the  trophozoite  of  forming  a  variable 

number  of  spores  in  an  individual. 
Monosporous. — The  character  of  the  trophozoite  of  forming  a  single  spore. 
Pansporoblast. — Coined  by  Gurley  (1893:408)  used  here  in  the  same  mean- 
ing, an  enclosed  area  in  the  endoplasm  of  the  vegetative  form,  in  which 

two  sporoblasts  become  dififerentiated. 


289]  STUDIES  ON  MYXOSPORIDIA—KUDO  51 

Plasmogamy.    Fusion  of  two  trophozoites,  coined  by  Doflein  (1898). 
Plasmotomy. — Division  of  trophozoite  into  daughter  individuals,  coined  by 

Doflein  (1898). 
Polar  capsule. — The  pyriform  or  spherical,  hollow  body  in  the  spore  which 

forms  a  polar  filament. 
Polar  filament. — The  filament  which  is  coiled  inside  the  polar  capsule. 
Polysporous. — The  character  of  the  trophozoite  of  forming  spores,  more 

than  two. 
Posterior  filament. — Fine  posterior  appendage  of  the  spore. 
Posterior  processes. — Posterior  differentiations  of  the  shell. 
Ridge. — Linear  or  network-like  elevation  of  the  shell  of  the  spore. 
Shell. — The  envelope  of  the  spore. 

Shell-valves. — Two  valves  which  compose  the  shell  of  the  spore. 
Sporoplasm. — The  protoplasmic  mass  found  inside  of  the  spore  (amebula 

or  sporozoite) ,  usually  situated  in  the  posterior  portion  of  the  spore. 
Sutural  diameter. — Same  as  length. 

Sutural  edge. — The  edge  of  the  shell-valves  cut  by  the  sutural  plane. 
Sutural  line. — The  line  on  the  shell  of  the  spore  marked  by  the  sutural  plane. 
Sutural  plane. — The  plane  on  which  two  shell- valves  meet  together. 
Sutural  ridge. — The  ridge  marking  the  sutural  line. 
Tail. — The  posterior  prolongation  of  the  valves  from  the  median  posterior 

end;  it  may  be  a  single  process  or  bifurcated. 
Thickness. — See  breadth. 

Trophozoite. — The  vegetative  or  multiplicative  stage  of  a  Myxosporidian. 
Vegetative  form. — Same  as  trophozoite. 


52 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[290 


CLASSIFICATION  OF  MYXOSPORIDIA 

The  classification  of  Myxosporidia,  was  first  carried  out  by  Thelohan 
as  early  as  1892,  who  considered  rightly  that  the  spore  was  the  only  reliable 
means  for  the  purpose.  In  1899  and  1901,  Doflein  introduced  into  the 
classification  two  Legions,  Disporea  and  Polysporea,  and  a  new  family. 
This  plan  has  generally  been  followed  by  various  authors  in  dealing  with 
these  protozoa.* 

The  classification  of  the  said  author,  however,  no  longer  agrees  with 
our  present  knowledge  of  the  animals.  In  the  first  place,  as  was  pointed 
out  by  some  authors,  for  instance  Davis  (1917:217),  it  is  far  from  being 
correct  to  divide  the  Myxosporidia  into  two  Legions,  Disporea  and  Poly- 
sporea, on  the  basis  of  the  number  of  spores  formed  in  each  vegetative 
form,  since  this  differs  even  in  one  and  the  same  species  as  was  observed 
by  Leger,  Auerbach,  Awerinzew,  Parisi,  Georgevitch,  Davis,  Kudo  and 
others  (see  Table  IV  on  page  53). 

Auerbach  who  had  observed  numerous  interesting  facts  in  this  group, 
had  adopted  Doflein's  classification  in  his  splendid  work  (1910)  by  simply 
adding  two  genera,  Zschokkella  and  Lentospora,  to  the  family  Myxidiidae. 
In  the  following  year  (1911),  he  tried  a  new  classification,  on  the  same 
basis  as  Doflein  did,  by  introducing  two  new  Legions  besides  these  two 
already  existing,  and  by  discarding  all  the  families.    Thus: 


I  Monosporea 
II  Mictosporea 


III  Disporea 
rv  Polysporea 


a)  Genus 

a)  Genus 

b)  Genus 

c)  Genus 

d)  Genus 

e)  Genus 

f)  Genus 

a)  Genus 

b)  Genus 

a)  Genus 

b)  Genus 

c)  Genus 

d)  Genus 

e)  Genus 


Coccomyxa 

Zschokkella 

Myxoproteus 

Myxidium 

Sphaeromyxa 

Chloromyxum 

Sphaerospora 

Ceratomyxa 

Leptotheca 

Myxosoma 

Lentospora 

Myxobolus 

Henneguya 

Hoferellus 


As  will  be  distinctly  seen  from  Table  IV,  the  classification  not  only  fails 
to  improve  Doflein's  classification  in  bringing  together  the  genera,  Myxo- 


*  Doflein  still  uses  the  same  classification  in  his  recent  work  (1916). 


291) 


STUDIES  ON  MYXOSPORIDIA—KUDO 


S3 


proteus,  Myxidium  and  Sphaeromyxa  into  Mictosporea,  and  Lentospora 
and  Henneguya  into  Polysporea,  but  increases  the  confusion  concerning 
relationship  among  the  genera. 


TABLE  IV 


Genus 


Mono- 

Mono- 

Mono-, 

and 

and 

Di- 

di-  and 

di- 

poly- 

sporous 

poly- 

sporous 

sporous 

sporous 

8 

3 

23 
2 

■■ 

1 

2 

2 

1 

2 
2 

1 

1 

2 

2 

1 

2 

1 
1 

Di-  and 

poly- 
sporous 


Poly- 
sporous 


Unknown 


Leptotheca 
15  species.... 

Ceratomyxa 
35  species.... 

Myxoproteus 
3  species 

Wardia* 

2  species 

Mitraspora* 

3  species 

Chloromyxum 

22  species... 
Sphaerospora 

10  species 

Sinuolinea* 

5  species 

Myxidium 

26  species 

Sphaeromyxa 

7  species , 

Zschokkella 

4  species 

M5Tcosoma 

3  species 

Lentosp)ora 

6  species 

Myxobolus 

63  species 

Henneguya 

32  species 

Hoferellus 

1  species 


1 
1 
4 

2 
1 
9 

5 
1 

3 

2 

54 

25 

1 


7 
5 
1 
1 

7 
2 

10 
2 
1 

2 
9 
5 


*  These  three  genera  are  unknown  to  Auerbach,  except  two  species  which  were  formerly 
placed  in  Leptotheca  and  Sphaerospora. 


54  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [292 

Parisi  (1912)  followed  Auerbach  in  his  paper  dealing  with  Myxosporidia 
from  Italian  waters.  Poche  (1913)  put  Auerbach's  classification  in  better 
form  as  follows: 

Order:  MjTcosporidia 

2  Superfamily  Mictosporea 

2  Family  Myxidiidae 

2  Genus    Zschokkella 

3  Genite    Myxoproteus 

4  Genus    Myxidium 

5  Genus    Sphaeromyxa 

6  Genus    Sphaerospora 

3  Family  Chloromyxidae 

7  Genus    Chloromyxum 

3  Superfamily  Disporea 

4  Family  Ceratomyxidae 

8  Genus    Ceratomyxa 

9  Genus    Leptotheca 

4  Superfamily  Polysporea 

5  Family  Myzosomatidae  (Poche) 

10  Genus    Myxosoma 

11  Genus    Lentospora 

6  Family  Myxobolidae 

12  Genus    Myxobolus 

13  Genus    Henneguya 

14  Genus    Hoferellus 


For  the  same  reason  given  in  discussing  Auerbach,  this,  however,  is  not 
conformable  with  the  present  state  of  knowledge  regarding  these  protozoa. 

It  was  not  until  1917  that  the  classification  of  the  Myxosporidia 
approached  to  a  more  natural  state  in  the  valuable  work  by  Davis  (1917: 
219-221).  He  pointed  out  sharply  the  unsatisfactory  features  in  Doflein's 
classification  and  proposed  a  different  system  as  follows: 

Order:  Myxosporidia. 

Suborder  I  Myxosporea  Davis 
Family  1  Ceratomyxidae 

Genus  1    Leptotheca 

Genus  2    CeratomjTca 
Family  2  Sphaerosporidae  Davis 

Genus  1    Myxoproteus 

Genus  2    Sphaerospora 

Genus  3    Sinuolinea 
Family  3  Myxidiidae 

Genus  1    Myxidiima 

Genus  2    Sphaeromyxa 

Genus  3    Zschokkella 
Family  4  Chloromyxidae 

Genus  1  Chloromyxum 


2931 


STUDIES  ON  MYXOSPORJDIA—KUDO 


55 


Suborder  n  Cystosporea  Davis 
Family  1  Myxosomidae*  Davis 
•  Genus  1    Myxosoma 

Genus  2    Lentospora 
Family  2  Myxobolidae 

Genus  1    Myxobolus  • 

Genus  2    Henneguya 
Genus  3    Hoferellus 

Thus,  Davis  selected  the  form  of  the  spore  for  the  establishment  of  two 
suborders  and  further  rearranged  the  genera  into  closer  positions  to  show 
relationship  to  each  other  better  than  any  one  of  the  previous  authors. 
He,  however,  named  the  suborders  according  to  a  secondary  character, 
i.e.,  the  seat  of  the  parasites  in  the  host.  According  to  his  definition  the 
trophozoites  of  the  species  belonging  to  Myxosporea  are  "with  few  excep- 
tions free  living  in  the  body-cavity,"  while  those  of  Cystosporea  "with 
few  exceptions"  are  tissue  parasites. 

From  TABLE  III  on  page  45,  are  taken  the  following  data  regarding 
this  point: 


Total  nimiber 

of  species 

known 


Number  of 

species  found  in 

body  cavity 


Number  of 

species  found  in 

tissue 


Number  of 

species  found  in 

both  places 


Seat 
unknown 


M)ntosporea. 
Cystosporea. 


132 
105 


114 

7 


14 

95 


Thus  it  appears  that  the  terms  Myxosporea  and  Cystosporea  do  not 
seem  to  be  properly  used.  These  may  be  replaced  by  terms  that  denote 
the  first  and  common  character  of  the  suborders. 

The  suggestions  as  to  the  adoption  of  other  characters  than  the  spore 
for  the  divisions  of  Myxosporidia,  proposed  by  Awerinzew  (1907:831; 
1908:64),  Auerbach  (1910:161)  and  Davis  (1917:217)  can  only  be  applied 
in  the  future.  At  the  present  time,  the  characters  concerning  the  vegetative 
form  do  not  appear  to  afford  a  better  and  more  natural  basis  for  the 
classification  of  Myxosporidia  than  those  of  the  spore.  Thus  from  the 
taxonomic  point  of  view  the  present  situation  does  not  seem  to  be  much 
improved  as  compared  with  that  at  the  end  of  the  last  century. 

The  writer  proposes  in  the  following  pages  a  new  classification  based  on 
the  characters  of  the  spore. 


*  Davis  did  not  notice  the  establishment  of  the  family  Myxosomatidae  by  F.  Poche  (1913), 
including  exactly  the  same  genera.    See  page  54. 


56  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [294 

Order  MYXOSPORIDIA  Butschli  1881 
Suborder  EURYSPOREA  nom.  nov. 
Largest  diameter  of  the  spore  at  right  angles  to  the  sutural  plane. 
One  polar  capsule  on  each  side  of  the  plane.  Sporoplasm  with  no  iodin- 
ophilous  vacuole.  Vegetative  form  found  in  body  cavity  (except  2  species) . 
Great  majority  parasites  of  marine  fish.  Monosporous,  disporous  and 
polysporous. 

Family     CERATOMYXIDAE  Dofiein     1899 
With  the  characters  of  the  suborder. 

Genus    LEPTOTHECA  Thelohan     1895 
Shell-valves  of  spore  hemispherical  or  shortly  rounded,      15  species. 
Disporous  (7  unknown).    14  species  in  body-cavity;  1  in  tissue;  all  in  marine 
fish.    Type  species:  Leptotheca  agilis  Thelohan. 

Genus  CERATOMYXA  Thelohan  1892 
Shell- valves,  conical  and  hollow,  attached  on  the  bases;  free  ends 
extended,  tapering  to  more  or  less  sharply  pointed  or  rounded  ends. 
Sporoplasm  usually  does  not  fill  the  cavity,  but  is  located  asymmetrically 
in  it.  35  species.  Disporous  (23  species),  monosporous  and  disporous 
(3  species),  disporous  and  polysporous  (4  species)  and  unknown  (5  species). 
All  (except  2  species  in  urinary  bladder)  in  the  gall-bladder  of  marine  fish. 
Type  species:  Ceratomyxa  arcuata  Thelohan. 

Genus     MYXOPROTEUS  Doflein  1898    emend.  Davis  1917 
Spores  roughly  pyramidal;  with  or  without  distinct  processes  from  the 
base  of  the  pyramid.    3  species.    Disporous  (one  species  unknown).    All 
in  urinary  bladder  of  marine  fish.    Type  species:  Myxoproteus  atnhiguus 
(Thelohan)  Doflein. 

Genus  WARDIA  nov.  gen. 
Spore  form  of  isosceles  triangle  with  two  convex  sides.  Oval  in  profile. 
Surface  of  shell  with  fine  ridges  which  turn  into  fringe-like  processes  at  the 
posterior  end.  The  polar  capsules,  large  and  perfectly  spherical,  situated 
at  the  central  portion  of  the  spore,  opening  at  the  anterior  tip.  Two  spe- 
cies. Polysporous  (one  species  unknown).  Tissue  parasite  (one  species) 
of  fresh-water  fish  and  amphibia,  both  found  in  Illinois,  U.  S.  A.  Type 
species:  Wardia  ovinocua  nov.  spec. 

Genus     MITRASPORA  Fujita  1912  emend.  Kudo 
Spores  spherical  or  ovoidal.    Two  polar  capsules  pyriform,  one  situated 
on  each  side  of  the  sutural  plane.     Shell  longitudinally  striated;  with  or 
without  long  and  fine  filaments  projecting  posteriorly  in  a  row  at  right 


295]  STUDIES  ON  MYXOSPORJDIA— KUDO  57 

angles  to  the  sutural  plane  at  the  posterior  side.  3  species.  Disporous 
and  polysporous.  All  found  in  kidney  of  fresh- water  fish.  Type  species: 
Mitraspora  cyprini  Fujita. 

Suborder     SPHAEROSPOREA    nom.  nov. 
Spores  spherical  or  subspherical,    with    two  'to   four   polar   capsules. 
Sporoplasm  without  iodinophilous  vacuole.     Vegetative  form  found  in 
body-cavity    and    tissue.      Monosporous,    disporous    and    polysporous. 
Parasites  of  marine  and  fresh-water  fish  and  amphibia. 

Family     CHLOROMYXIDAE  Thelohan     1892* 

Spores  with  four  polar  capsules.  Monosporous,  disporous  and  poly- 
sporous. 

Genus     CHLOROMYXUM  Mingazzini     1890 

With  the  characters  of  the  family.  22  species.  18  in  body  cavity; 
4  in  tissue.  7  from  marine  and  12  from  fresh-water  fish,  2  in  amphibia, 
1  in  insect.    Type  species:  Chloromyxutn  leydigi  Mingazzini. 

Family     SPHAEROSPORIDAE  Davis     1917 
Spores  with  two  polar  capsules.    Monosporous,  disporous  and  polyspo- 
rous. 

Genus     SPHAEROSPORA  Thelohan     1892 
Spores  with  two  polar  capsules.     Monosporous,  disporous  and  poly- 
sporous.   10  species.    Body-cavity  and  tissue.    5  from  fresh-water  and  5 
marine  fish.    Type  species:  Sphaerospora  divergens  Thelohan. 

Genus     SINUOLINEA  Davis     1917 
Spores  with  or  without  lateral  processes.    Two  polar  capsules  spherical. 
Sutural  line  sinuous.     5  species.     Disporous  and  polysporous.     In  the 
urinary  bladder  of  marine  fish.    Type  species:  Sinuolinea  dimorpha  Davis. 

Suborder     PLATYSPOREA     nom.  nov. 
Sutural  plane  of  the  spore  coincides  with  or  at  an  acute  angle  to  the 
longest  diameter.    One  or  two  polar  capsules.    Sporoplasm  with  or  without 
an  iodinophilous  vacuole. 

Family     MYXIDIIDAE  Thelohan     1892 
Two  polar  capsules,  one    at    each    end.     Sporoplasm   without  any 
iodinophilous  vacuole.    Spores  fusiform. 


*  Th61ohan  (1892)  used  the  terms:  Chloromyxidees,  Myxidid^es,  Myxobolid^es,  which 
Gurley  (1893)  made  over  into  Chloromyxidae,  Myxidiidae,  Mj^obolidae,  so  that  the  credit 
of  recognizing  and  establishing  these  families  belongs  to  Thelohan. 


58  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [296 

Genus  MYXIDIUM  ButschU  1882 
Spores  more  or  less  regularly  fusiform,  with  pointed  or  rounded  ends. 
Polar  filaments  long  and  fine.  26  species.  Monosporous,  disporous  and 
polysporous.  22  in  body-cavity;  4  in  tissue.  15  in  marine  and  6  in  fresh- 
water fish,  2  in  fishes  from  both  waters  and  3  in  reptilia.  Type  species: 
Myxidium  Ueberkuhni  Biitschli. 

Genus    SPHAEROMYXA  Thelohan     1892 
Spores  fusiform,  with  truncated  ends.    Polar  filament  short  and  thick. 
Trophozoites  large  and  disc  shaped.    7  species.    Polysporous  (2  unknown). 
6  in  body-cavity;  1  unknown.    6  in  marine  fish;  1  in  amphibia.     Type 
species:  Sphaeromyxa  halhianii  Thelohan. 

Genus  ZSCHOKKELLA  Auerbach  1910 
Spores,  semicircular  in  front  view;  pointed  at  ends.  Polar  capsules 
large  and  spherical,  opening  on  the  flat  edge  near  the  tips.  Sutural  line 
usually  curved  in  S-form.  4  species.  Monosporous,  disporous  and  poly- 
sporous. Body-cavity.  2  from  marine  and  2  from  fresh-water  fish.  Type 
species:  Zschokkella  hildae  Auerbach. 

Family    MYXOSOMATIDAE  Poche     1913 
Two  polar  capsules  at  the  anterior  end.    Sporoplasm  without  iodino- 
philous  vacuole. 

Genus    MYXOSOMA  Thelohan     1892 
Spores  ovoidal,  flattened  and  more  or  less  elongated.    3  species.    Poly- 
sporous.   Tissue  parasites.    2  in  fresh-water  and  1  in  marine  fish.    Type 
species:  Myxosoma  dujardini  Thelohan. 

Genus    LENTOSPORA  Plehn     1905 
Spores  similar  to  Myxobolus  in  form.  Sporoplasm  without  any  iodino- 
philous  vacuole.    6  species.    Disporous  and  polysporous  (2  unknown).    1 
in  marine  and  3  in  fresh-water  fish,  2  from  fishes  in  both  waters.    Type 
species:  Lentospora  cerebralis  (Hofer)  Plehn. 

Family  MYXOBOLIDAE  Thelohan     1892 
Spores  with  one  or  two  polar  capsules  at  the  anterior  end,  with  or 
without  posterior  processes.     Sporoplasm  with  an  iodinophilous  vacuole. 
Majority  polysporous  in  fresh-water  fishes. 

Genus  MYXOBOLUS  Biitschli  1882 
Spores  ovoidal  or  ellipsoidal;  flattened.  One  or  two  polar  capsules  at 
the  anterior  end.  Shell  without  posterior  process.  63  species.  Poly- 
sporous (9  species  unknown).  59  species  in  tissue;  4  unknown.  5  in 
marine  and  56  in  fresh-water  fish,  1  in  annelid  and  1  in  amphibia.  Type 
species:  Myxobolus  mUlleri  Biitschli. 


2971  STUDIES  ON  MYXOSPORIDIA—KUDO  59 

Genus  HENNEGUYA  Thelohan  1892 
Spores  more  or  less  globular  or  ovoidal.  Two  polar  capsules  at  the 
anterior  end.  Posterior  end  of  the  shell-valves  prolonged  into  more  or  less 
extended  processes,  which  unite  and  form  a  tail  in  the  median  line.  32 
species.  Polysporous,  disporous  and  monosporous.  28  species  in  tissue 
and  4  in  body-cavity.  In  fresh- water  fish,  except  one.  Type  species: 
Henneguya  psorospertnica  Thelohan. 

Genus    HOFERELLUS  Berg     1898 
Spores  pyramidal,  with  two  posterior  processes  from  the  lateral  faces. 
1  species.     Polysporous.     Tissue  and  body-cavity  of  fresh-water  fish. 
Type  and  only  species:  Hoferellus  cy print  Doflein. 


eO  '  ILLINOIS  BIOLOGICAL  MONOGRAPHS '  [298 


DESCRIPTIONS  OF  GENERA  AND  SPECIES 
Suborder  EURYSPOREA  nom.  nov. 
The  definition  of  the  suborder  is  recorded  on  page  56. 

Family  CERATOMYXIDAE  Doflein 
1899        Ceratomyxidea  Doflein  1899  :  378 

1901        Ceratomyxidae  Doflein  1901  :  182 

The  characters  of  the  family  are  described  on  page  56, 

Genus  LEPTOTHECA  Thelohan 
1895        Leptotheca  Thelohan  1895  :  331 

The  characters  of  the  genus  are  described  on  page  56. 
Type  species:  Leptotheca  agilis  Thelohan. 

LEPTOTHECA  AGILIS  Thelohan 
[Figs.  1  to  5] 
1892        Ceratomyxa  agilis  Th61ohan  1892  :  962 

1895        Leptotheca  agilis  Thflohan  1895  :  332 

1898        Leptotheca  agilis  Doflein  1898  :  294, 297 

Habitat:  Gall-bladder  of  Trygon  pastinaca  L.  and  Scorpaeona  sp.; 
France,  Rovigno,  Napoli. 

Vegetative  form:  Form  generally  elongated.  Anterior  end  rounded 
where  a  mass  of  fat  globules  is  found,  while  the  posterior  end  terminates 
in  a  point.  Size  not  exceeding  85^1  by  20  to  25/n.  The  posterior  part  is 
sometimes  divided  into  a  certain  number  of  lobes.  In  the  protoplasm, 
the  globules  are  clearly  seen.  Pseudopodia  are  localized  at  the  anterior 
portion  of  the  body.  They  are  long,  40  to  50/i  in  length,  filiform  and  very 
active  in  moving  from  back  toward  front,  just  like  the  motion  of  oars. 
Disporous. 

Spore:  Slightly  elongated.  Dimensions:  sutural  diameter  6  to  7n, 
breadth  11  to  12/t. 

LEPTOTHECA  ELONGATA  Thelohan 
[Fig.  6] 

1895        Leptotheca  dongata  Thflohan  1895  :  332 

1898        Leptotheca  dongata  Doflein  1898  :  312 

1917        Leptotheca  dongata  Georgevitch         1917b  :  99-106 

Habitat:  Gallbladder  of  Merluccius  merluccius  L.  (M.  vulgaris)  and 
Motdla  tricirrata;  Marseille,  Banyuls,  Le  Croisic,  Napoli,  Monaco. 


2991  STUDIES  ON  MYXOSPORIDIA—KUDO  61 

Vegetative  form:  Form  variable.  Many  individuals  show,  however, 
a  very  characteristic  form.  It  is  elongated  and  has  the  length  of  about 
120/i.  The  anterior  end  is  enlarged  into  a  disc-shaped  depression,  on  the 
edge  of  which,  the  branched  pseudopodia  are  formed.  The  body  gradually 
narrows  itself  toward  the  posterior  end.  Also  club-shaped,  etc.  The  short 
lobose  pseudopodia  show  no  movement  like  that  of  oars. 

Georgevitch's  form:  Young  forms,  oval  or  rounded,  are  attached  to 
the  epithelial  cells  of  the  bladder  with  a  long  filiform  pseudopodium  at  the 
free  end.    Such  forms  often  agglomerate  in  great  number. 

Spore:  Form  similar  to  the  spore  of  Leptotheca  agilis.  Dimensions  on 
the  average:  Sutural  diameter  12  to  IS/x,  breadth  18  to  20/x. 

LEPTOTHECA  POLYMORPHA  (Thelohan)  Labbe 
1895        Leptotheca  elongata  Thelohan  1895  :  332-333 

1899        Leptotheca  polymorpJia  Labb6  1899  :  88 

Habitat:  Gall-bladder  of  Phycis  mediterraneus  {P.  phycis  L.);  Banyuls. 

Vegetative  form:  Form  extremely  polymorphous,  with  three  main 
types.  1)  Somewhat  regularly  club-shaped,  with  lobose  pseudopodi^,,  some- 
times filiform  at  one  end.  2)  Irregular  as  is  the  case  with  Ceratomyxa 
truncata,  with  long  (ISfx)  ectoplasmic  processes,  which  are  motionless  or 
very  slow  in  motion.  Lobose  pseudopodia  are  formed  actively.  3)  More 
or  less  rounded  with  bristle-like  filose  pseudopodia.  Intermediary  forms 
are  also  found.  Often  many  individuals  unite  together.  The  protoplasm 
is  much  different  from  other  forms,  i.e.,  more  homogeneous  and  compact. 
Granules  are  hardly  visible  on  account  of  vacuolar  appearance. 

Spore:  Dimensions:  sutural  diameter  10  to  12)ti,  breadth  18  to  20jti, 
length  of  polar  filament  40m. 

LEPTOTHECA  PARVA  Thelohan 
[Fig.  7] 
1895        Leptotheca  parva  Thelohan  1895  :  333 

1912        Leptotheca  parva  Auerbach  1912  :  42-43 

Habitat:  Gall-bladder  of  Scomber  scombrus  L.;  Marseille,  Le  Croisic, 
Le  Vivier-sur-mer,  Kristiansund,  Stavanger,  Bergen. 

Vegetative  form:  Form  ordinarily  rounded,  spherical  or  subspherical. 
Often  club-shaped.  Size  not  larger  than  12  to  15/x  in  diameter.  Proto- 
plasm finely  granular.    Pseudopodia  lobose. 

Spore:  Small,  more  or  less  elongated,  curved  in  arch-form.  Dimen- 
sions: sutural  diameter  3  to  4jLt,  breadth  8  to  IOai. 

LEPTOTHECA  RENICOLA  Thelohan 
1895        Leptotheca  renicola  Thelohan  1895:333 

Habitat:  Urinary  tubules  of  the  kidney  of  Scomber  scombrus  L.;  Mar- 
seille, Le  Croisic. 


62 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[300 


Vegetative  form:  Small.    No  marked  character. 

Spore:  Globular.   Form  similar  to  the  spore  of  Sphaerospora.    Dimen- 
sions: sutural  diameter  Sn,  breadth  10/x. 


LEPTOTHECA  HEPSETI  Thelohan 
1895        LepMkeca  hepseti  Th€lohan  1895  :  334 

Habitat:  Gall-bladder  of  Atherina  hepsetus  L.;  Marseille.  Of  rare 
occurrence;  Thelohan  observed  it  but  once. 

Vegetative  form:  Not  described. 

Spore:  Form  triangular  with  rounded  angles.  Dimensions:  sutural 
diameter  7  to  Sju,  breadth  12  to  15/i. 

LEPTOTHECA  PERLATA  (Gurley)  Labbe 
[Fig.  8] 
1883  Balbiani  1883  :  201,  204 

1894        CUoromyxum  {Sphaerospora)  perlatutn    Gurley  1894  :  272 
1899        LepMheca  perlata  Labb6  1899  :  88 

Habitat:  Acerina  cernua  L.;  France  (?). 
Vegetative  form:  Not  described. 

Spore:  Elliptic.  Two  small  polar  capsules  converging.  Dimensions 
not  given. 

LEPTOTHECA  sp.  Awerinzew 
[Figs.  16, 17] 

1908  LepMheca  sp.  Awerinzew  1908  :  51, 52 

Habitat:  Gall-bladder  of  Sehastes  norvegicus;  Eastern  Finmark? 
Vegetative  form:  Rounded  form  with  clear  differentiation  of  proto- 
plasm into  ectoplasm  and  endoplasm.    Plasmotomy  occurs. 
Spore:  Undescribed.    No  figure. 

LEPTOTHECA  MACROSPORA  Auerbach 
[Fig.  9] 

1909  LepMheca  macrospora  Auerbach  1909  :  70-71 

1910  LepMheca  macrospora  Auerbach  1910b  :  768-769 
1910        LepMheca  macrospora              Auerbach  1910c  :  167 
1912        LepMheca  macrospora               Auerbach  1912  :  42-43 

Habitat:  Gall-bladder  of  Sehastes  viviparus  H.  KT.ai,ndS.dactylopterus; 
Bergen,  Kristiansund  (May,  September). 

Vegetative  form:  Trophozoites  spherical  with  the  average  diameter 
of  26  to  30/1.  Homogeneous  ectoplasm  layer  exhibits  somewhat  active 
ameboid  movements.  Endoplasm,  granular  in  living  specimen,  is  rather 
sharply  distinguishable  from  the  ectoplasm  and  contains  large  nuclei. 


301]  STUDIES  ON  MYXOSPORIDIA—KUDO  63 

Spore :  Size  large.  Form  resembles  to  that  of  Leptotheca  parva.  Dimen- 
sions: sutural  diameter  and  tliickness  I3n,  breadth  26/i.  Polar  capsules 
short  oval,  with  a  length  of  5.2/i,  length  of  polar  filament  about  130m 
(KOH).  In  the  second  host,  a  few  normal  and  numerous  abnormal  spores 
with  three  or  four  polar  capsules  were  observed. 

LEPTOTHECA  INFORMIS  Auerbach 
[Fig.  10] 

1910        Leptotheca  informis  Auerbach  1910b  :  770-771 

1912        Leptotheca  informis  Auerbach  1912  :  42-44 

Habitat:  Gall-bladder  of  Molva  vulgaris  Flem.  and  Gadus  merlangus; 
Bergen,  Tjomo. 

Vegetative  form:  Young  trophozoites  with  somewhat  long  and  narrow 
pseudopodia  formed  of  hyaline  ectoplasm;  movements  active.  The  proto- 
plasm is  differentiated  into  ectoplasm  and  endoplasm.  When  stained,  two 
large  (7  to  9/i)  and  two  small  (3  to  4n)  nuclei  were  observed  in  an  individual, 
27m  long  excluding  the  pseudopodia.  Sporulating  trophozoites  are  gen- 
erally round  and  each  forms  two  spores,  which  are  developed  independently 
to  each  other  (i.e.,  not  in  ordinary  pansporoblast).  Auerbach  observed 
centrosomes  in  the  nuclei  of  larger  type  in  division.    Disporous. 

Spore:  Large  and  heavily  built.  Greatly  curved.  Sutural  line  fairly 
well  marked.  Polar  capsules  round.  Dimensions:  sutural  diameter  lO/t, 
breadth  18  to  20m,  thickness  9m,  diameter  of  polar  capsules  3  to  4m.  Sporo- 
plasm  contains  two  nuclei,  3.5  to  4m  in  diameter. 

LEPTOTHECA  LONGIPES  Auerbach 
[Fig.  11] 
1910        Leptotheca  longipes  Auerbach  1910b  :  771 

1912        Leptotheca  longipes  Auerbach  1912  :  42^3 

Habitat:  Gall-bladder  of  Brosmius  brosme  Asc;  Bergen  (May). 

Vegetative  form:  Trophozoites  elongated  or  rounded.  Only  few 
pseudopodia  which  are  very  long.  Small  forms  with  a  very  long  process, 
were  observed  in  large  numbers;  length  of  the  body  being  10m,  while  the 
process  was  60m  long.  Endoplasm  contains  nuclei  of  various  sizes.  Disporous. 

Spore:  Form  similar  to  that  of  Leptotheca  informis,  though  smaller. 
Dimensions:  sutural  diameter  8  to  9m,  breadth  12  to  14m,  thickness  8m, 
diameter  of  polar  capsule  2.5m. 

LEPTOTHECA  FUSIFORMIS  Davis 
[Fig.  12] 
1917        Leptotheca  fiisiformis  Davis  1917:222 

Habitat:  Gall-bladder  of  Cestracion  zygaena;  Beaufort  (July). 


64  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [302 

Vegetative  form:  Pyriform,  tapering  gradually  toward  the  posterior 
end,  which  usually  terminates  in  a  long,  slender  process;  colorless  and 
transparent.  Progressive  movements  rapid.  Endoplasm  granular,  the 
granules  being  more  abundant  at  the  anterior  end.  The  average  size  of 
full-grown  individuals:  50/i  by  13/i.     Disporous. 

Spore:  Elliptical  in  front  view;  fusiform  in  side  view.  Sutural  plane 
slightly  oblique  to  the  longest  diameter,  the  line  forming  a  marked  ridge. 
Polar  capsules  open  on  opposite  sides  of  the  spore.  Sporoplasm  finely 
granular,  confined  to  the  central  part  of  spore.  Dimensions :  sutural  diame- 
ter 9/i,  breadth  16)u,  polar  capsule  4.5m  long,  length  of  polar  filament  30/i. 

LEPTOTHECA  SCISSURA  Davis 
[Fig.  13] 
1917        Leptotheca  scissura  Davis  1917  :  222 

Habitat:  Gall-bladder  of  Dasybatis  hastatus  and  Pteroplatea  maclura 
Le  Sue;  Beaufort  (July,  August). 

Vegetative  form:  Young  form  elongated,  with  long  attenuated  posterior 
process;  usually  sUghtly  constricted  just  posterior  to  rounded  anterior  end, 
which  bears  numerous  long,  filiform  pseudopodia.  Progressive  movement 
rapid.  Ectoplasm  distinguishable  at  the  anterior  end.  Endoplasm  usually 
filled  with  small,  clear,  colorless  spherules,  which  become  larger  and 
yellowish  as  the  body  increases  in  size.  Each  spherule  contains  one  to 
several  dark-brown  granules,  which  increase  in  size  and  number  and  finally 
collect  in  an  irregular  clump  at  the  centre  of  spherule.  The  larger  indi- 
viduals are  usually  flattened  dorso-ventrally.  The  posterior  end  is  divided 
into  long  slender  processes,  presenting  sometimes  a  network  caused  by  the 
fusion  of  two  or  more  adjacent  processes.  Full-grown  forms:  length  125  to 
150/i,  breadth  20  to  25^.    The  longest  observed,  195^1  by  16jLt.    Disporous. 

Spore:  Elliptical  in  front  view;  somewhat  flattened  along  the  posterior 
side.  Sutural  line  distinct  and  at  right  angles  to  the  longest  diameter. 
Polar  capsules  have  foramina  at  some  distance  from  the  capsular  margin. 
Sporoplasm  finely  granular,  nearly  filling  both  valves.  Dimensions: 
sutural  diameter  ll/x,  breadth  22/li,  diameter  of  polar  capsule  4^. 

LEPTOTHECA  LOBOSA  Davis 
[Fig.  14] 
1917        Leptotheca  lobosa  Davis  1917  :  223 

Habitat:  Urinary  bladder  of  Paralichthys  dentatus  L.;  Beaufort  (July). 

Vegetative  form:  Usually  spherical  which  may  form  a  large  rounded 
pseudopodium  composed  of  ectoplasm.  Body  colorless  and  transparent 
to  translucent.  Ameboid  movements  very  slow.  Ectoplasm  contains 
coarse  granules,  which  are  of  uniform  size  and  very  distinct.    Endoplasm 


303]  STUDIES  ON  MYXOSPORIDIA— KUDO  65 

less  granular  and  more  transparent  than  ectoplasm,  containing  numerous 
large,  yellow,  fat  globules,  which  are  abundant  in  large  forms.  Diameter 
up  to  24ai.    Disporous. 

Spore:  Elliptical  in  front  view;  valves  slightly  tapering  but  rarely 
alike.  Sutural  line  forming  a  sinuous  ridge.  Polar  capsules  open  at  some 
distance  from  the  anterior  margin.  Sporoplasm  nearly  filling  both  valves. 
Free  spores  are  often  seen  to  remain  united  at  the  sutural  line.  Dimensions: 
sutural  diameter  9  to  10/i,  breadth  16  to  IS/u;  diameter  of  polar  capsule  3/t. 

LEPTOTHECA  GLOMEROSA  Davis 
[Fig.  15] 
1917        Leptotheca  glomerosa  Davis  1917  :  223 

Habitat:  Urinary  bladder  of  Faralichthys  albiguttus]  Beaufort. 

Vegetative  form:  Rounded  or  somewhat  irregular  in  shape,  with  short 
iobose  pseudopodia.  Body  transparent  and  colorless.  Ameboid  move- 
ments slow.  Ectoplasm  hyaline,  forming  a  distinct  outer  layer.  Endo- 
plasm  finely  granular,  with  numerous  small  fat  globules  varying  in  size. 
Almost  entire  body  is  used  for  spore  formation.  Diameter  of  rounded 
sporulating  trophozoite  about  llfx.    Disporous. 

Spore:  Approximately  cylindrical;  valves  rounded  at  ends.  The  coiled 
polar  filament  not  visible  in  the  polar  capsule.  Sutural  line  at  right  angles 
to  the  longest  diameter.  Sporoplasm  finely  granular,  fills  the  extracapsular 
cavity  of  spore.  Dimensions:  sutural  diameter  4.5/i,  breadth  9/u,  diameter 
of  polar  capsule  2/t. 

Genus  CERATOMYXA  Thelohan 

1892        Ceratomyxa  Thdohan  1892  :  169,  171,  175 

1895        Ceratomyxa  Thelohan  1895  :  334 

The  characters  of  the  genus  described  on  page  56. 
Type  species:  Ceratomyxa  arcuata  Thelohan. 

CERATOMYXA  ARCUATA  Thelohan 


[Figs. 

18  to  22] 

1892 

Ceratomyxa  arctuita 

Thelohan 

1892a 

:1091 

1895 

Ceratomyxa  arcuata 

Thelohan 

1895  : 

335-336 

1899 

Ceratomyxa  arcuata 

Labb6 

1899  : 

90 

1912 

Ceratomyxa  arcuata 

Parisi 

1912  : 

290-291 

1913 

Ceratomyxa  arcuata 

Jameson 

1913  : 

2 

1916 

Ceratomyxa  arcuata 

Georgevitch 

1916a 

:3 

Habitat:  Gall-bladder  of  Pagellus  centrodontus  C.  et  V.,  Crenilabrus 
melops  L.,  Motella  tricirrata  Bl.,  Ophidium  vasalli,  Gohius  paganellus  L., 
Heliases  chromis  Gthr.;  Scot paena  scrofa  L.,  S.  porcus  L.;  France  (Marseille, 
Banyuls,  Concarneau,  Roscoff),  Italy  (Napoli,  summer),  Monaco  (May). 


66  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [304 

Vegetative  form:  Polymorphous;  generally  club-shape,  pseudopodia 
localized  at  the  broad  end;  the  other  end  cylindrical  or  terminating  in  a 
sharp  point.  Some  other  different  forms.  Pseudopodia,  always  localized, 
lobose  pointed  at  the  extremities.  Ectoplasm  hyaline  and  thin.  Endo- 
plasm  contains  fat  globules  and  particular  elements,  mostly  large  refractive 
globules,  which  seem  to  disappear  in  the  sporulating  individuals.  Dimen- 
sions (maximum):  length  35  to  40/x,  breadth  12  to  IS/i,  pseudopodia  about 
10m  loiig-    Disporous. 

Spore:  Arch  form.  Shell  valves  equal.  Sporoplasm  occupies  the 
extracapsular  cavity  of  the  spore.  The  length  varies  rather  considerably 
according  to  the  development  of  the  lateral  processes,  which  are  occasionally 
acuminated  or  very  short.  Often  extremities  are  rounded.  Dimensions 
(Thelohan) ;  breadth  20  to  30/i,  sutural  diameter  5  to  8/i.  Parisi's  measure- 
ments: breadth  25  to  31/i,  sutural  diameter  5.5  to  6/i,  length  of  polar 
capsules  3.5  to  An,  length  of  polar  filaments  25/x. 

Remarks:  The  writer  agrees  with  Parisi  in  eliminating  Labbe's  two 
subspecies  (1899:90),  as  they  are  too  arbitrary. 

CERATOMYXA  SPHAERULOSA  Thelohan 


[Figs. 

23  to  24] 

1892 

Ceratomyxa  sphaerulosa 

Thelohan 

1892 

171 

1895 

Ceratomyxa  sphaerulosa 

Thelohan 

1895 

334-335 

1909 

?  Ceratomyxa  sphaerulosa 

Auerbach 

1909 

80 

1912 

Ceratomyxa  sphaerulosa 

Auerbach 

1912 

4,45 

1916 

Ceratomyxa  sphaerulosa 

Georg^vitch 

1916a 

:3 

Habitat:  Gall-bladder  of  Mustelus  canis  Mitch.  (M.  vulgaris),  Galeus 
galeus  L.  (G.  canis),  Clupea  harengus,  Scullium  canicula  Cuv.;  St-Valery- 
en-Caux,  Roscoff,  Bergen,  Monaco  (May). 

Vegetative  form:  Form  more  or  less  definite  among  the  adults.  Gen- 
erally elongated.  Both  ends  slightly  attenuated.  Wide  in  the  middle  part 
of  the  body.  Lobose  pseudopodia  at  one  of  the  extremities.  Others 
more  massive  or  more  or  less  regularly  spherical,  in  which  case  the  pseu- 
dopodia are  formed  from  the  whole  surface.  Spherical  form  does  not 
exceed  50  to  60iu  in  diameter.  Other  forms  90  to  lOO/x  by  30  to  40/i  (largest). 
Young  forms  colorless  and  are  more  variable  than  the  adults.  Protoplasm 
homogeneous  and  finely  granular.  Adult  form,  on  the  contrary,  yellowish 
or  greenish  yellow.  The  endoplasm  is  filled  with  small  (5ju  in  diameter) 
spheres,  in  the  centre  of  which  5  to  6  small  granules,  yellowish  brown  or 
greenish  in  color,  are  present.    Disporous. 

Spore:  Remarkably  large.  Polar  filament  can  be  seen  in  vivo  (easily 
extruded  by  KOH,  ether,  etc.)  Sporoplasm  occupies  one  of  the  shell- valves, 
while  a  small  mass  of  very  pale  looking  substance  is  seen  in  the  other. 
Dimensions:  sutural  diameter  10  to  12/i,  breadth  90  to  100/x,  subspherical 
polar  capsule  6  to  7  by  5/i,  sporoplasm  12  to  15  by  8  to  9/i. 


305]  STUDIES  ON  MYXOSPORIDIA—KUDO  67 

CERATOMYXA  PALLIDA  Thelohan 


1895 

Ceratomyxa  pallida 

TWlohan 

1895  :  336-337 

1898 

Ceratomyxa  pallida 

Doflein 

1898  :  341 

1916 

Ceratomyxa  pallida 

Georg6vitch 

1916b  :  2,  3 

Habitat:  Gall-bladder  of  Box  boops  L.  and  B,  salpa  L.;  Marseille, 
Villefranche,  Rovigno,  Monaco  (May). 

Vegetative  form:  Ordinarily  spherical  not  exceeding  16  to  20/i  in 
diameter.  Many  individuals  often  found  in  massive  groups.  Pseudopodia 
lobose  and  mostly  short.    Protoplasm  extremely  pale  with  fine  granules. 

Spore:  Dimensions:  sutural  diameter  5^,  breadth  25  to  30)u. 

CERATOMYXA  GLOBULIFERA  Thelohan 
[Fig.  25] 
1895        Ceratomyxa  globulifera  Th61ohan  1895  :  338 

Habitat:  Gall-bladder  of  Merluccius  merluccius  L.  {M.  vulgaris) \ 
Marseille,  Banyuls. 

Vegetative  form:  Polymorphous.  Elongated  into  long  branches, 
including  endoplasm.    Endoplasm  contains  small  refractive  globules. 

Spore:  Elongated.  Shell- valves  unequal,  one  being  longer  and  finer 
than  the  other.    Dimensions:  sutural  diameter  10/i,  breadth  50/x. 

CERATOMYXA  APPENDICULATA  Thelohan 

[Fig.  26] 

1892        Ceratomyxa  appendiculata         Th61ohan  1892a  :  963-964 

1895        Ceratomyxa  appendiculata         Thelohan  1895  :  337 

1898        Ceratomyxa  appendiculata         Doflein  1898  :  300, 311 

Habitat:  Gall-bladder  of  Lophius  piscatorius  L.,  L.  budegassa  Spin.; 
RoscoflF,  Le  Croisic,  Marseille,  Banyuls,  Napoli,  Rovigno. 

Vegetative  form:  Extremely  polymorphous.  Young  form  spherical, 
spatulaform,  club-shape,  etc.  In  adult  form,  the  main  thick  part  of  the 
body,  in  which  spore  formation  takes  place,  forms  1  to  6  long  prolongations, 
twice  or  three  times  longer  than  the  main  part  of  the  body.  Pseudopodia 
lobose,  filiform  or  elongated  with  enlargements.    Disporous. 

Spore:  Lateral  prolongations  of  shell- valves  well  developed.  Dimen- 
sions: sutural  diameter  5  to  J/x,  breadth  50/i. 

CERATOMYXA  TRUNCATA  Thelohan 
[Fig.  27] 

1895        Ceratomyxa  iruncata  Th61ohan  1895  :  336 

1912        Ceratomyxa  truncata  Parisi  1912  :  289-290 

Habitat:  Gall-bladder  of  Clupea  pilchardus  Walb.  {Alosa  sardina); 
Marseille,  Villefranche,  Napoli  (August,  September). 


68  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [306 

Vegetative  form:  Polymorphous.  Ordinarily  more  or  less  rounded, 
with  lobose  pseudopodia.  Pseudopodia  long  and  often  shows  very  active 
movements.  Endoplasm  very  finely  granular,  contains  small  fat  globules 
which  are  found  in  irregular  mass  or  in  a  circular  form.    Disporous. 

Spore:  Valves  are  short  and  truncate.  Sporoplasm  occupies  the  whole 
cavity.  Spores  with  three  valves  are  frequently  encountered.  Dimensions : 
breadth  25jLt,  5^  in  sutural  diameter.  According  to  Parisi,  spores  with  two 
shell- valves  are  rather  few  (10%),  while  those  with  three  (70%)  and  four 
shell- valves  (20%)  prevail  in  number!  Dimensions:  breadth  20  to  30/*, 
length  of  the  polar  filament  45ac. 


CERATOMYXA  RETICULARIS  Thelohan 

[Fig.  28] 

1895        Ceratomyxa  reticularis  Th61ohan  1895  :  337-338 

Habitat:  Gall-bladder  of  Trachinus  draco  L.;  Banyuls. 

Vegetative  form:  Extremely  polymorphous.  Generally  spherical  or 
club-shaped.  Well  developed  trophozoites  have  the  similar  form  as  in 
C.  appendiculata.    Endoplasm  highly  reticular,  with  refringent  fluid. 

Spore:  Shell  valves  are  short  and  truncate,  one  of  which  is  curved  to  the 
rear.    Dimensions:  sutural  diameter  12  to  15/x,  breadth  45  to  50/i. 

CERATOMYXA  INAEQUALIS  Doflein 

[Fig.  29] 

1898        Ceratomyxa  inaequalis  Doflein  1898  :  284-285 

Habitat:  Gall-bladder  of  Crenilabrus  mediterraneus  and  C.  pavo; 
Napoli. 

Vegetative  form:  Form  usually  club-shaped.  Protoplasm  in  active 
motion,  is  differentiated  distinctly  into  ectoplasm  and  endoplasm.  Body 
yellowish  brown  by  the  presence  of  granules  in  endoplasm.  Inactive  forma- 
tion of  pseudopodia.  Ameboid  movements  or  progressive  movements  by 
means  of  the  posterior  process.  Size:  20  to  40m  by  5  to  10/x  in  average. 
Length  of  the  posterior  process  up  to  30/i.  After  spore  formation,  only 
two  nuclei  remain  in  protoplasm,  which  apparently  degenerate  later. 
Disporous. 

Spore:  Elliptical,  somewhat  flattened.  Massive.  Very  transparent. 
Both  ends  round,  but  unequally  built,  i.e.,  one  end  is  club-shaped.  Polar 
capsules  are  somewhat  round  and  are  bound  to  the  shell  by  protoplasmic 
bridges.  The  polar  filament  is  not  seen  in  fresh  spores.  Dimensions: 
sutural  diameter  6ju,  breadth  31/i,  diameter  of  the  polar  capsule  2.5  to 
3fi,  length  of  polar  filament  is  half  breadth  of  the  spore  (diluted  nitric  acid). 


307]  STUDIES  ON  MYXOSPORJDIA— KUDO  69 

CERATOMYXA  LINOSPORA  Doflein 
[Figs.  30  to  31] 
1898        Ceralomyxa  Unospora  Doflein  1898  :  285 

Habitat:  Gall-bladder  of  Labrus  turdus;  Napoli. 

Vegetative  form:  Club-  or  spindleshape.  Protoplasm  highly  granu- 
lated. Body  whitish  grey,  though  very  transparent.  Pseudopodia  very 
fine  and  only  formed  at  the  anterior  end  of  the  body.  Size:  30  to  35jw  by 
16  to  18/x.    Disporous. 

Spore:  Form  symmetrical  with  long  thread-like  lateral  processes.  In 
sporoblast,  the  processes  are  wound  around  the  spore.  It  is  twice  as  long  as 
the  breadth  of  the  spore.  Polar  capsules  large  and  spherical  pyriform. 
Dimensions:  total  breadth  50^,  breadth  of  the  main  part  of  the  spore  10 
to  12fi,  sutural  diameter  5ju,  length  of  lateral  process  20/i.  "Polar  filament 
was  too  fine  to  be  measured." 

CERATOMYXA  RAMOSA  Awerinzew 

[Figs.  32  and  33] 

1907        Ceralomyxa  ramosa  Awerinzew  1907  :  831-834 

*1908        Ceralomyxa  ramosa  Awerinzew  1908  :  60-66 

Habitat:  Gall-bladder  oi  Hippoglossus  vulgaris  Flemm.;  Kjellebjord, 
Murman  coast. 

Vegetative  form:  Form  irregular  ameboid,  owing  to  the  presence  of 
peculiar  pseudopodia.  The  middle  part  of  the  body  is  enlarged  into  an 
ellipsoidal  form,  where  nuclei  and  sporoblasts  are  present.  From  this  part 
two,  rarely  one  or  three  processes  are  formed,  which  branch  out  several 
pseudopodia  of  different  length.  The  finer  portions  of  pseudopodia  anasto- 
mose each  other  and  form  a  characteristic  and  remarkable  network. 
Differentiation  of  protoplasm  is  not  very  distinct.  Ectoplasm  is  not  well 
developed,  tho  covering  the  entire  surface  of  the  body  as  a  thin  la)'er. 
Endoplasm  slightly  vacuolated  and  granular,  forms  the  greater  part  of  the 
body.    Disporous  and  polysporous. 

Spore:  Form  and  size  (?)  resemble  C.  arcuata.  Slightly  curved  toward 
the  posterior  side.  Valves  usually  unequally  built,  one  being  longer  than 
the  other.  Sporoplasm  almost  always  asymmetrically  situated  in  the  shell. 
Polar  capsules  on  each  side  of  the  sutural  plane  and  of  the  plane  perpen- 
dicular to  the  sutural  plane,  cutting  the  spore  into  two  equal  parts.  Young 
spores  in  development  ellipsoidal  to  kidne)^  bean  shape.  Dimensions: 
sutural  diameter  12  to  20/i,  breadth  50  to  80/x. 

*  Professor  J.  Zeitlin  has  kindly  translated  some  part  of  the  paper,  for  which  the  writer 
expresses  his  thanks. 


70  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [308 

CERATOMYXA  DREPANOPSETTAE  Awerinzew 
[Figs.  34  to  39] 

1907  Ceratomyxa  sp.  Awerinzew  1907  :  832-833 

1908  Ceratomyxa  drepanopsettae        Awerinzew  1908  :  1-41, 45-47 

1909  Ceratomyxa  drepanopsettae        Awerinzew  1909:74-112 

1912  Ceratomyxa  drepanopsettae        Auerbach  1912  :  44-45 
1918        Ceratomyxa  drepanopsettae        Kudo  1918  :  14-15 

Habitat:  Gall-bladder  of  Pleuronectes  platessa,  P.flesus,  Drepanopsetta 
plaUessoides,  Hippoglossus  vulgaris,  Hippoglossoides  limandoides  and 
Paralichihys  deniatus;  Murmankuste,  Kabelvaag,  Rorvik,  Tjomo,  Woods 
Hole  (August,  September). 

Vegetative  form:  Polymorphous.  Usually  very  much  elongated  and 
slender  forms.  Protoplasm  differentiated.  Endoplasm  coarsely  granular. 
Pseudopodia  lobose  and  filiform  (2  to  3n),  with  which  the  trophoz9ites 
attach  themselves  to  the  epithelium  of  the  bladder.    Disporous. 

Spore:  Curved  toward  the  posterior  side.  Shell  with  rounded  ends. 
Valves  almost  always  unequally  built.  Dimensions:  breadth  50  to  80/i. 
Auerbach's  form:  Form  variable.  Size:  sutural  diameter  about  12  to 
14/*,  breadth  about  56/i,  diameter  of  polar  capsule  about  4  to  6/x,  length  of 
the  cavity  in  which  the  sporoplasm  is  located  about  34n.  Kudo's  form: 
Variable.  Sutural  diameter  8  to  10m,  average  breadth  64^,  diameter  of 
polar  capsule  6/i. 

CERATOMYXA  TYLOSURI  Awerinzew 
[Figs.  40  and  41] 

1913  Ceratomyxa  tylosuri  Awerinzew  1913a  :  153 

Habitat:  Gall-bladder  of  Tylosurus  schismaiorhynchus;  Lorengo 
Marques,  Delagoa  Bay  (Africa). 

Vegetative  form:  Large,  irregular,  disc-like  or  large  ameboid,  with 
blunt  lobose  pseudopodia  and  highly  granular  protoplasm. 

Spore:  Large.  The  anterior  edge  arch-shape,  while  the  posterior  edge 
has  two  small  horns  which  are  located  symmetrically  to  the  sutural  line. 
Polar  capsules  elongated  and  are  separated  from  binuclear  sporoplasm  by  a 
special  membrane.  Rarely  spore  with  three  polar  capsules.  Dimensions 
breadth  124  to  140^,  sutural  diameter  40  to  45/*,  thickness  25  to  30/«. 

CER.\TOMYXA  (?)  SPARI  Awerinzew 

[Figs.  42  and  43] 

1913        Ceratomyxa  (?)  spari  Awerinzew  1913a :  153-154 

Habitat:  Gall-bladder  of  Sparus  berda;  Lorenfo  Marques,  Delagoa 
Bay  (Africa). 


309]  STUDIES  ON  MYXOSPORIDIA—KUDO  71 

Vegetative  form:  Large  (100  to  120/*),  disc-form  ameboid,  containing 
a  large  number  of  enclosures  and  granules  of  different  size.  In  one  case,  a 
number  of  this  form,  carrying  no  spore,  underwent  budding,  which  resulted 
in  forming  spherical  forms  of  various  size,  some  of  which  divided  again  into 
2  to  6  parts  (Plasmotomy?).    Monosporous  and  disporous. 

Spore:  More  or  less  curved.  Two  polar  capsules  lie  closely  together 
on  each  side  of  the  sutural  plane.  Ends  of  shell- valves  are  rounded. 
Dimensions:  breadth  50  to  60/x,  sutural  diameter  12  to  IS/it,  thickness 
12  to  15ai,  polar  filament  very  long  (length  not  given). 

Remarks:  Awerinzew  thinks  this  is  the  intermediate  form  between 
Leptotheca  and  Ceratomyxa. 

CERATOMYXA  sp.  (?)  Awerinzew 
1913        Ceratomyxa  sp.  (?)  Awerinzew  1913a  :  154 

Habitat:   Gall-bladder  of  Scatophagus  argus;  Delagoa  Bay  (Africa). 

Vegetative  form:  Small,  disc-form  ameboid  (25  to  35/i),  containing 
two  spores  of  indistinct  contour,  on  account  of  incomplete  formation  of  the 
shell.  Two  spores,  apparently,  developed  in  one  pansporoblast.  Dis- 
porous. 

Spore:  Form  could  not  exactly  be  made  out.  Polar  capsules  were 
arranged  like  those  of  other  Ceratomyxa. 

CERATOMYXA  sp.  (?)  Awerinzew 
1913        Ceratomyxa  sp.  (?)  Awerinzew  1913a  :  154-155 

Habitat:  Gall-bladder  of  Rhinobathus  Awer.  (?);  Lorenzo  Marques 
(Africa). 

Vegetative  form:  Irregular  shape.  Endoplasm  highly  granular.  In  the 
epithelial  layer  of  the  gall-bladder  numerous,  spherical  cysts  (30  to  35ju) 
were  found.    Two  spores  are  formed  in  one  pansporoblast.    Disporous. 

Spore:  Cylindrical  with  broad  and  slightly  rounded  ends.  Dimensions: 
sutural  diameter  16  to  19/x,  breadth  70  to  80/*,  thickness  16  to  19ju. 

CERATOMYXA  ACADIENSIS  Mavor 
[Figs.  44  to  47] 

1915  Ceratomyxa  acadiensis  Mavor  1915  :  27-30 

1916  Ceratomyxa  acadiensis  Mavor  1916  :  551-574 

Habitat:  Gall-bladder  of  Urophycis  chuss  (trophozoites  are  attached  to 
undetermined  Myxosporidia,  see  p.  176),  Zoarces  angularis,  Pseudo- 
pleuronectes  americanus ;  New  Brunswick  (Canada)  (July  to  September). 

Vegetative  form:  Polymorphous.  Typically  club-shaped  with  very 
long  tail,  or  irregularly  stellate.    Pseudopodia  show  rigidity.    Sometimes 


72  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [310 

clumps  of  protoplasm  along  their  length,  which  are  connected  by  thin 
hyaline  filaments  of  ectoplasm.  Differentiation  of  protoplasm  is  usually 
observable  at  the  anterior  region.  Dimensions:  length,  excluding  tail, 
12  to  15^1,  breadth  10  to  20/x,  tail  up  to  60/i.    Disporous. 

Spore:  Wide,  short  and  slightly  compressed  dorso-ventrally,  with  very 
long  fine  lateral  filaments.  Polar  capsules  spherical.  Polar  filament 
invisible  in  vivo.  Dimensions:  breadth  40  to  50/i,  sutural  diameter 
7  to  8m,  diameter  of  polar  capsule  3  to  4^,  length  of  polar  filament  70^, 
length  of  lateral  filaments  250  to  300/x. 

CERATOMYXA  sp.  Georgevitch 
1916        Ceratomyxa  sp.  Georgevitch       1916a  :  3 

Habitat:  Gall-bladder  of  Muraena  sp.;  Monaco  (May). 
Vegetative  form:  No  description. 
Spore:  No  description.    No  figure. 

CERATOMYXA  CORIS  Georgevitch 
[Fig.  48] 

1916  Ceratomyxa  coris  Georgevitch         1916a  :  4-5 

1917  Ceratomyxa  coris  Georgevitch         1917a  :  1-20 

Habitat:  Gall-bladder  of  Coris  julius,  C.  giofredi;  Villefranche  (March, 
June). 

Vegetative  form:  Various  forms,  club-shape,  spherical  or  elongated, 
with  lobose  or  filiform  pseudopodia.    Disporous  and  rarely  Polysporous. 

Spore:  More  or  less  ellipsoidal.  Lateral  prolongations  of  the  shell- 
valves  short  and  truncate.  Sutural  line  straight.  Sporoplasm,  elongate, 
rounded,  elliptical,  fills  a  part  of  the  extracapsular  cavity  of  the  spore. 
Polar  capsules  rounded,  almost  spherical,  not  converging.  Dimensions 
not  given. 

Remarks:  Georgevitch  observed  (1917:  Fig.  30)  that  spores  of  Glugea 
marionis  occurred  in  disporous  trophozoite  of  Ceratomyxa  coris,  which  he 
thought  to  have  happened  accidentally  by  plasmogamy  of  these  two  Cnido- 
sporidia.  The  above  mentioned  figure,  however,  strongly  suggests  that 
G.  marionis  may  be  leading  parasitic  life  in  the  trophozoite  of  C.  coris. 

CERATOMYXA  HEROUARDI  Georgevitch 
[Fig.  49] 

1913  Leptotheca  (?)  sp.  Jameson  1913  : 2 

1916  Ceratomyxa  herouardi  Georgevitch  1916a  :  5-8 

1916  Ceratomyxa  herouardi  Georgevitch  1916b  :  717-19, 983-985 

1917  Ceratomyxa  herouardi  Georgevitch  1917  :  375-399 

Habitat:  Gall-bladder  of  Box  Salpa  L.;   Monaco  (May). 


3111  STUDIES  ON  MYXOSPORIDIA— KUDO  73 

Vegetative  form:  Polymorphous.  Elongated  with  same  breadth  or 
tapering  to  one  end;  club-shaped  with  roundish  enlargements.  Young 
trophozoites  spherical  or  pyriform.  Pseudopodia  long  and  narrow  or 
broad  and  bi-  or  multi-lobate.  Body  colorless  both  in  the  young  and  the 
adult.  Protoplasm  homogeneous  and  finely  granular.  Disporous  and 
polysporous.  Spores  are  found  inside  of  the  endoplasm  and  in  the  roundish 
buds,  ordinarily  two  spores  being  formed  in  each  bud.  Number  of  buds 
on  one  trophozoite  varies.    Plasmotomy  by  budding  and  division. 

Spore:  Elongated  elliptic.  Polar  capsules  spherical  and  large.  Sutural 
plane  cuts  the  spore  into  exactly  equal  two  parts.  Two  nuclei  in  sporoplasm 
are  rather  small  and  are  always  in  one  of  the  shell-valves.  Dimensions 
not  given. 

Remark:  The  form,  mentioned  by  Jameson  in  the  same  seat,  host  and 
locality,  that  "has  something  of  the  appearance  of  a  Leptotheca"  and  that 
is  also  "almost  certainly  neither  of  the  two  Myxosporidia — Ceratomyxa 
pallida  Sind  Henneguy a  neapolitana  .  .  .  ,"  is  probably  identical  with  the 
present  form. 

CERATOMYXA  MESOSPORA  Davis 

[Fig.  50] 

1917        Ceratomyxa  mesospora  Davis  1917  :  223-224 

Habitat:  Ga,\\-h\a.dder  oi  Cestracion  zygaena,  C .  tiburo;  Beaufort  (July). 

Vegetative  form:  Pyriform,  elongate,  with  long,  slender  posterior 
process.  Numerous  filiform  pseudopodia  at  anterior  end.  Progressive 
movements  rapid.  Body  colorless.  No  sharp  demarcation  between 
ectoplasm  and  endoplasm.  Endoplasm  finely  granular  and  filled  with 
small,  colorless,  homogeneous  spherules.  Spherules  absent  at  anterior  end. 
Size:  total  length  70  to  85^,  length  exclusive  of  posterior  process  50  to  75ju, 
breadth  20  to  25/1.    Disporous. 

Spore:  Greatly  elongate,  each  valve  forming  a  slightly  tapering  cone, 
rounded  at  the  apex.  Valves  not  compressed.  Sutural  plane  forming  an 
acute  angle  with  the  longest  diameter.  Polar  capsules  conspicuous.  Coiled 
polar  filaments  very  distinct.  Polar  capsules  are  remarkable  in  that  they 
are  asymmetrically  situated,  one  being  always  located  in  the  widest  part 
of  the  spore,  while  the  other  being  a  little  to  one  side.  Sporoplasm  asym- 
metrically situated,  sometimes  being  entirely  confined  to  the  larger  valve. 
Dimensions:  breadth  50  to  65/i,  sutural  diameter  about  8^,  diameter  of 
polar  capsule  4.5/x,  length  of  polar  filament  90jLt. 

Remarks:  Similar  to  C.  sphaerulosa  Thel.  and  occurs  with  C.  recurvata 
Davis  in  the  same  organ. 

CERATOMYXA  SPHAIROPHORA  Davis 

[Fig.  51] 
1917        Ceratomyxa  sphairophora  Davis  1917  :  224 

Habitat:  Gall-bladder  of  Scoliodon  terrae-novae;  Beaufort. 


74  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [312 

Vegetative  form  rPyriform,  elongate.  Numerous  fine  filiform  pseudo- 
podia  at  anterior  end.  Progressive  movements  rapid.  Body  colorless  and 
transparent.  Ectoplasm  clear  and  homogeneous.  Structure  of  endoplasm 
highly  variable,  in  majority  of  trophozoites  filled  with  transparent  homo- 
geneous spherules.  Small  fat  globules  at  the  anterior  end.  In  some  sporu- 
lating  individuals,  the  endoplasm  shows  vacuolated  structure  without  any 
spherules,  usually,  however,  sporulating  trophozoites  exhibit  well-defined 
spherules.  The  spherules  or  vacuoles,  as  the  case  may  be,  are  separated  by 
a  thin  layer  of  distinctly  granular  endoplasm  containing  numerous  rod- 
shaped  or  rounded,  colorless  bodies,  which  in  their  appearance  are  strikingly 
like  small  bacteria  tho  they  are  not  bacteria,  as  they  fail  to  take  Giemsa 
stain.     Size  of  sporulating  trophozoites  100  to  llO/i  by  25/i.     Disporous. 

Spore:  Shell- valves  greatly  elongated,  tapering  gradually  toward  the 
ends.  Long,  attenuated  ends  of  valves  hollow  and  so  fragile  that  it  is 
almost  impossible  to  find  an  example  in  which  they  are  not  more  or  less 
distorted.  Sutural  plane  perpendicular  or  only  slightly  oblique  to  the 
longest  diameter.  Polar  capsules  are  spherical  and  large;  slightly  conver- 
gent, opening  some  distance  apart  on  the  anterior  side.  Coiled  polar 
filament  distinct.  Sporoplasm  confined  to  large,  central  part  of  spores, 
but  extending  farther  into  one  valve  than  the  other.  Dimensions:  total 
breadth  115  to  140/:,  sutural  diameter  about  12/i,  diameter  of  polar  capsules 
(>H,  length  of  polar  filament  75/z. 

CERATOMYXA  TAENIA  Davis 

[Figs.  52  and  53] 
1917        CeraUmyxa  taenia  Davis  1917  :  224-225 

Habitat:  Gall-bladder  of  ScoUodon  terrae-novae;  Beaufort. 

Vegetative  form:  Similar  to  those  of  C.  sphairophora  Davis,  and  no 
character  has  been  found  by  which  they  may  be  distinguished.  Sporulating 
trophozoites  can  be  easily  distinguished  on  account  of  the  very  different 
appearance  of  the  spore  and  their  different  arrangement  within  the  tropho- 
zoites. The  spores  of  this  species  are  situated,  as  is  usually  the  case  in 
Ceratomyxa,  with  the  greater  part  of  the  spore  parallel  to  the  long  axis  of 
the  trophozoite,  only  a  part  of  one  valve  being  bent  back  along  the  rest 
of  the  spore.  Size:  sporulating  trophozoites  length  SO/x,  breadth  25/i. 
Disporous. 

Spore:  Valves  greatly  elongated.  Shell  very  thin,  the  membrance  on 
opposite  sides  of  each  valve  being  in  contact  for  about  two-thirds  of  its 
length,  forming  a  thin  ribbonlike  structure;  basal  third  of  each  valve  only 
slightly  compressed;  terminal  ribbonlike  portion  of  each  valve  usually 
twisted  so  that  plane  of  ribbon  is  at  right  angles  to  the  main  part  of  Jjje 
spore.  Polar  capsules  small,  pyriform  to  spherical  and  convergent.  Coiled 
polar  filament  indistinct.     Sutural  plane  perpendicular  to  the  longest 


3131  STUDIES  ON  MYXOSPORIDIA—KUDO  75 

diameter.  Sporoplasm  finely  granular,  filling  the  basal  third  of  each  valve, 
sometimes  extending  farther  into  one  valve  than  the  other.  Dimensions: 
breadth  140  to  150/i,  breadth  of  central  portion  45jli,  sutural  diameter  6/Lt, 
diameter  of  the  polar  capsules  3/t. 

CERATOMYXA  ATTENUATA  Davis 

[Fig.  54] 

1917        Ceratomyxa  attenuata  Davis  1917  :  225 

Habitat:  Gall-bladder  of  ScoUodon  terrae-novae;  Beaufort  (July). 

Vegetative  form:  Elongate,  pyriform,  with  long,  tapering  posterior 
process;  at  anterior  end  numerous  long  filiform  pseudopodia.  Progressive 
movements  rapid.  Ectoplasm  distinct  only  at  anterior  end.  Endoplasm 
filled  with  small,  refractive,  yellowish  or  brownish  granules,  which  are 
uniformly  distributed  throughout  the  trophozoite.  Between  the  brownish 
granules,  the  endoplasm  is  clear  and  colorless,  not  granular,  except  at 
extreme  anterior  end  where  it  contains  a  clump  of  small  fat  globules.  Size 
of  full-grown  trophozoites  100  to  120  by  27  /u.    Disporous. 

Spore:  Valves  greatly  elongated;  a  symmetrical,  one  valve  being  about 
15/i  shorter  than  the  other  and  ending  abruptly;  the  longer  valve  tapering 
gradually  to  a  point.  About  midway  of  each  valve,  is  a  thin  septum; 
external  to  the  septum  the  valves  are  empty.  Polar  capsules  are  large, 
opening  on  the  anterior  margin.  Coiled  polar  filaments  distinct.  Sutural 
plane  oblique  to  longitudinal  axis,  usually  forming  a  ridge.  Sporoplasm 
asymmetrically  situated  in  central  part  of  the  spore.  Dimensions:  breadth 
llSju,  sutural  diameter  9n,  diameter  of  polar  capsules  4.5/1,  length  of  polar 
filament  60jLt. 

CERATOMYXA  RECURVATA  Davis 

[Figs.  55  and  56] 

1917        Ceratomyxa  recurvata  Davis  1917  :  225-226 

Habitat:  Gall-bladder  of  Cestracion  zygaena;  Beaufort  (July). 

Vegetative  form:  Pyriform  with  long,  slender  posterior  process. 
Body  colorless.  Actively  motile,  forming  filiform  pseudopodia  of  ectoplasm 
at  anterior  end.  Endoplasm  colorless  and  granular,  filled  with  large, 
homogeneous  spherules.  Full-grown  trophozoites  130  to  175)u,  length  of 
the  main  body  about  lOOfi.  Spores  are  developed  singly  from  distinct 
sporoplasts  and  not  necessarily  in  pairs.  Disporous  and  polysporous  (up 
to  10  spores,  6  and  8  are  common  numbers). 

Spore:  Valves  greatly  curved  toward  the  posterior  side,  usually  sym- 
metrical, but  occasionally  one  may  be  much  more  incurved  than  the  other. 
Valves  circular  in  cross  section  at  the  base  but  toward  the  ends  greatly 


76  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [314 

flattened.  Ends  of  valves  sharply  pointed.  Polar  capsules  large,  opening 
at  some  distance  from  the  anterior  margin.  Coiled  polar  filaments  dis- 
tinct. Sporoplasm  finely  granular  usually  extending  farther  into  one  valve 
than  the  other.  Dimensions:  breadth  between  points  of  greatest  curva- 
ture about  16/11,  sutural  diameter  8  to  9/i,  diameter  of  polar  capsules  4.5/*. 

CERATOMYXA  LUNATA  Davis 

[Figs.  57  to  60] 

1917        CeraUmyxa  lunata  Davis  1917  :  226-227 

Habitat:  Gall-bladder  of  Galeocerda  tigrinus;  Beaufort  (August). 

Vegetative  form:  Pyriform,  rounded  after  being  on  the  slide  for  some 
time.  Progressive  movements  slow  Endoplasm  filled  with  large,  homo- 
geneous spherules,  which  are  usually  colorless,  sometimes  light  yellow. 
At  extreme  anterior  end,  the  endoplasm  contains  numerous  small  fat 
globules.     Disporous. 

Spore:  Considerably  variable  in  size  and  form.  The  larger  and  more 
typical  are  more  or  less  crescent-shaped;  symmetrical;  valves  curved 
toward  rear,  terminating  in  more  or  less  rounded  ends.  Polar  capsules 
large  and  open  on  opposite  sides  of  spore.  Coiled  polar  filament  distinct. 
Sporoplasm  finely  granular,  symmetrically  situated  in  spore.  Smaller 
spores  differ  from  large  ones  chiefly  in  size;  valves  are  much  shortened 
and  have  a  greater  curvature,  with  more  distinctly  rounded  ends.  Dimen- 
sions: breadth  30m  (longest  38^), sutural  diameter  9/i,  diameter  of  polar 
capsules  4/*,  length  of  polar  filament  37/*.  Small  forms:  breadth  15/*, 
sutural  diameter  7/*,  diameter  of  polar  capsules  ^n. 

CERATOMYXA  ABBREVIATA  Davis 
[Fig.  61] 
1917        Ceratomyxa  abbreviala  Davis  1917  :  227 

Habitat:  Gall-bladder  of  Scoliodon  terrae-novae;  Beaufort  (August). 

Vegetative  form:  Elongate,  pyriform,  with  usually  a  very  long,  slender 
posterior  process.  Body  colorless.  Progressive  movements  rapid.  Dis- 
tinct differentiation  of  protoplasm,  posterior  process  usually  composed  of 
ectoplasm  (rarely  endoplasm  may  extend  into  it  for  a  short  distance). 
Pseudopodia,  short,  tapering  or  filiform  at  anterior  end.  Dimensions: 
length  up  to  90/*,  breadth  10  to  12/*,  diameter  of  rounded  sporulating 
trophozoites  about  27/t.    Disporous. 

Spore:  Roughly  crescent-shaped;  sutural  diameter  exceptionally 
great  in  comparison  with  the  breadth.  Ends  of  valves  ^rounded,  slightly 
asymmetrical.  Shell  exceptionally  tough  and  resistant  to  reagents.  Polar 
capsules  large,  prominent  and  open  on  opposite  sides  of  spore.  Sporoplasm 
finely  granular,  confined  entirely  to  one  valve.  Dimensions:  breadth  17/*, 
sutural  diameter  14/*,  diameter  of  polar  capsules  4.5/*. 


315]  STUDIES  ON  MYXOSPORIDI A— KUDO  77 

CER\TOMYXA  FLAGELLIFERA  Davis 
[Fig.  62] 
1917        Ceratomyxa  flageUifera  Davis  1917:227 

Habitat:  Gall-bladder  of  Carcharhinus  sp?;  Beaufort  (July). 

Vegetative  form:  Pyriform,  short,  tapering  toward  the  posterior  end, 
sometimes  dividing  into  a  number  of  long,  slender,  transparent  processes. 
Extremely  long  filiform  pseudopodia,  developed  at  anterior  end,  can  be 
seen  to  sweep  slowly  back  like  a  whiplash  until  they  come  to  lie  by  the  side 
of  the  body.  Progressive  movements  slow.  Ectoplasm  clear,  transparent, 
forming  a  distinct  layer  at  anterior  end.  Endoplasm  in  large  trophozoites 
filled  with  large  numbers  of  rod-shaped,  bacteria-like  bodies,  which  are  more 
abundant  in  the  anterior  half  than  in  the  posterior.  Endoplasm  in  younger 
trophozoites,  with  much  less  or  without  any  bacteria-like  bodies,  shows  a 
vacuolated  structure.  Size  up  to  115  to  I20fi  in  length  and  40  to  45/*  in 
breadth.     Disporous. 

Spore:  Valves  greatly  elongated,  conical,  with  rounded  ends.  Sutural 
ridge  well  marked.  Polar  capsules  large,  opening  on  opposite  sides*  of 
spore.  Coiled  polar  filament  very  distinct.  Sporoplasm  granular,  symmet- 
rically situated,  but  extending  only  a  short  distance  into  each  valve. 
Dimensions:  breadth  II8/1,  sutural  diameter  12/x,  diameter  of  polar  cap- 
sules 6/i. 

CERATOMYXA  AGGLOMERATA  Davis 

[Fig.  63] 

1917        Ceratomyxa  agglomerata  Davis  1917  :  228 

Habitat:  Gall-bladder  of  Synodus  foetans;  Beaufort. 

Vegetative  form:  Pyriform,  usually  with  long,  slender,  posterior 
process.  Body  colorless  and  transparent.  Actively  motile,  moving  by 
means  of  characteristic  wavelike  movements  of  the  ectoplasm,  from  which 
are  projected  numerous  short,  conical  to  filiform  pseudopodia.  Pseudo- 
podia travel  back  along  sides  of  body  for  about  one-third  its  length  and 
then  disappear,  new  ones  being  continually  formed  at  the  anterior  end. 
Ectoplasm  distinguishable  at  anterior  end.  Endoplasm  clear,  very  trans- 
parent, usually  homogeneous,  sometimes  finely  granular.  Large  numbers 
of  fat  globules  usually  present.  Size  of  sporulating  trophozoites  38/i  by 
12/i.     Disporous. 

Spore:  Asymmetrical,  one  valve  being  smaller  and  more  attenuated 
than  the  other;  larger  valve  compressed.  Polar  capsules  spherical.  Coiled 
polar  filaments  indistinct.  Sporoplasm  filling  nearly  entire  smaller  valve, 
but  only  extending  a  short  distance  into  the  larger  one.  Dimensions: 
breadth  24  to  28ju,  sutural  diameter  5ju,  diameter  of  polar  capsules  3ju. 


78  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [316 

CERATOMYXA  AMORPHA  Davis 
[Fig.  64] 
1917        Ceratomyxa  atnorpha  Davis  1917  :  228 

Habitat:  Gall-bladder  of  Synodus  foetans;  Beaufort. 

Vegetative  form:  Rounded  or  irregular  in  shape,  with  short  lobose 
pseudopodia;  not  pyriform;  slowly  ameboid.  Body  colorless.  Ectoplasm 
well  developed,  forming  a  distinct  layer;  transparent,  finely  granular. 
Endoplasm  granular,  with  large  numbers  of  small  fat  globules  scattered 
through  it  or  aggregated  into  one  or  two  large  clumps  (difference  between 
the  present  form  and  C.  agglomerata).    Disporous. 

Spore:  Asymmetrical;  crescent-shaped;  valves  short,  conical,  some- 
what compressed.  One  valve  distinctly  smaller  and  more  conical  than  the 
other.  Sutural  ridge  perpendicular  to  longitudinal  axis.  Polar  capsules 
large,  opening  at  some  distance  from  the  anterior  side.  Coiled  polar 
filaments  distinct.  Sporoplasm  granular,  asymmetrically  situated,  being 
chiefly  confined  to  smaller  valve.  Dimensions:  breadth  27/z,  sutural 
diameter  11/i,  diameter  of  polar  capsules  4ju. 

CERATOMYXA  MONOSPORA  Davis 

[Figs.  65  to  67] 

1917        Ceratomyxa  monospora  Davis  1917  :  228-229 

Habitat:  GdJi[-\Adididitr  oi  Peprilus  alepidotus;  Beaufort.  Abundantly 
present  in  June,  much  less  in  July,  being  entirely  absent  in  the  bladder  at 
the  end  of  the  month. 

Vegetative  form:  Pyriform,  with  a  slender  posterior  process  and  one  to 
several  filiform  pseudopodia  at  anterior  end.  Body  colorless  and  trans- 
parent. Movements  very  slow.  No  clear  differentiation  between  ecto- 
plasm and  endoplasm,  the  entire  body  being  composed  of  a  clear,  finely 
granular  protoplasm.  Fat  globules  more  abundant  in  larger  individual, 
which  are  aggregated  into  small  clumps.  Size  of  vegetative  trophozoites 
up  to  24/x  in  length  and  15m  in  width.  Monosporous  form  much  smaller 
than  disporous.  Monosporous  and  disporous.  Nearly  entire  substance 
of  trophozoite  is  used  up  in  spore  formation. 

Spore:  Crescent-shaped.  Valves  cylindrical,  tapering  toward  the  end, 
which  is  rounded  and  compressed.  Curvature  of  valves  varies.  One 
valve  is  more  attenuated  than  the  other.  Sutural  ridge  perpendicular  to 
the  longest  diameter.  Polar  capsules  large.  Sporoplasm  usually  asym- 
metrically situated.  Dimensions:  breadth  18  to  25/x,  sutural  diameter 
5  to  6jLi,  diameter  of  polar  capsules  3/i. 

Remarks:  This  species  is  evidently  very  close  to  C.  pallida  Th61. 
Similar  form  was  found  in  Prionotus  evolans  (gall-bladder),  which  showed 
somewhat  larger  trophozoites  and  spores  than  C.  monospora. 


3171  STUDIES  ON  MYXOSPORIDJA—KUDO  79 

CERATOMYXA  STREPTOSPORA  Davis 
[Figs.  68  and  69] 
1917        CeraUmyxa  streptospora  Davis  1917  :  229 

Habitat:  Gall-bladder  of  Chaetodipterus  faber;  Beaufort  (June,  but 
not  in  July). 

Vegetative  form:  Pyriform,  colorless  and  transparent.  A  few  conical, 
filiform,  wavelike  pseudopodia  at  anterior  end.  Ectoplasm  recognizable  at 
anterior  end.  Endoplasm  finely  granular,  with  a  few,  small,  fat  globules, 
filled  with  transparent,  homogeneous  spherules.  Size:  48  by  12/x  to  60 
by  9iJL.     Disporous. 

Spore:  Compressed  valves  greatly  elongated,  with  rounded  ends. 
Sutural  ridge.  Polar  capsules  spherical.  Coiled  polar  filament  indistinct. 
Sporoplasm  finely  granular,  entirely  filling  both  valves.  Dimensions: 
breadth  34  to  39/x,  sutural  diameter  4)u,  diameter  of  polar  capsules  3/*. 

CERATOMYXA  AGGREGATA  Davis 
[Fig.  70] 
•    1917        Ceratomyxa  aggregata  Davis  1917  :  229 

Habitat:  Gall-bladder  of  Leostomus  xanthurus,  Micropogon  undulatus; 
Beaufort  (July). 

Vegetative  form:  Form  rounded  to  somewhat  irregular  in  shape,  rarely 
pyriform;  slowly  ameboid.  Body  colorless  and  transparent.  No  clear 
differentiation  of  protoplasm.  Endoplasm  finely  granular,  containing 
numbers  of  small  fat  globules.  Sporulating  trophozoites  show  a  tendency 
to  collect  in  groups  composed  of  a  large  number  of  individuals  so  closely 
associated  that  it  is  often  impossible  to  make  out  the  individual  outlines. 
Size  of  full-grown  form  18/i  by  14/i.    Disporous. 

Spore:  Crescent-shaped;  valves  much  elongated,  tapering  toward  the 
ends,  which  are  compressed.  Polar  capsules  spherical  and  opaque.  Sporo- 
plasm granular,  situated  symmetrically  in  the  spore  cavity.  Dimensions: 
breadth  about  50jtt,  sutural  diameter  6  to  7/x,  diameter  of  polar  capsule 
3.5ai. 

CERATOMYXA  UNDUE  ATA  Davis 

[Fig.  71] 
1917        Ceratomyxa  undulata  Davis  1917  :  230 

Habitat:  Gall-bladder  of  Ancylopsetta  quadrocellata  Gill.;  Beaufort 
(June  to  August). 

Vegetative  form:  Pyriform,  sometimes  fusiform,  tapering  toward 
posterior  end.  Movements  rapid.  Body  colorless.  Ectoplasm  observable 
at  anterior  part,  constantly  undergoes  rapid,  wavelike  undulating  move- 
ments and  extrudes  fine  conical  or  filiform  pseudopodia.     Pseudopodia 


80  JLUNOIS  BIOLOGICAL  MONOGRAPHS  [318 

are  formed  very  rapidly  and  vary  in  length.  After  reaching  a  considerable 
length  the  pseudopodia  usually  travel  posteriorly  along  sides  of  body 
and  then  disappear.  Endoplasm  very  transparent,  often  vacuolated, 
containing  numerous  small  fat  globules.  Size  of  full-grown  trophozoite: 
25/1  by  10  to  12/i  in  average.     Disporous. 

Spore:  Crescent-shaped.  Valves  cylindrical,  not  compressed,  ends 
rounded,  one  valve  being  somewhat  longer  and  more  conical  than  the  other. 
Polar  capsules  convergent.    Coiled  polar  filaments  distinct.     Sporoplasm  i 

granular,  asymmetrically  situated,  sometimes  being  almost  confined  to 
more  conical  valve.  Dimensions:  breadth  22  to  24m,  sutural  diameter  6/i, 
diameter  of  polar  capsules  3/x. 

CERATOMYXA  NAVICULARIA  Davis 

[Figs.  72  and  73] 
1917        Ceratomyxa  navicularia  Davis  1917  :  230 

Habitat:  Urinary  bladder  of  Paralichthys  dentatus,  P.  albiguttus, 
Sphaeroides  maculatus;  Beaufort  (June  to  August). 

Vegetative  form:  Rounded  or  slightly  irregular  in  shape,  never  pyri- 
form.  Body  colorless.  Very  slow  ameboid.  No  distinct  ectoplasm. 
Entire  trophozoite  finely  granular,  containing  a  few  small  fat  globules. 
Nearly  entire  body  is  used  up  in  the  formation  of  spores.  Diameter  about 
17/1.     Disporous. 

Spore:  Variable  in  shape  and  size.  Symmetrical  or  asymmetrical, 
often  boat-shaped,  slightly  compressed  dorso-ventrally,  with  rounded 
ends.  Polar  capsules  convergent,  shows  polar  filament  indistinctly. 
Sporoplasm  finely  granular,  extending  into  both  valves,  but  usually 
somewhat  farther  into  one  than  the  other.  Dimensions:  breadth  14  to  22/x 
(average  16/Lt),  sutural  diameter  5  to  7.5m  (average  6m),  diameter  of  polar 
capsules  2m. 

CERATOMYXA  SPINOSA  Davis 

[Fig.  74] 

1917        Ceratomyxa  spinosa  Davis  1917  :  230 

Habitat:    Urinary  bladder  of  Paralichthys  albiguttus;  Beaufort. 

Vegetative  form:  Rounded  or  slightly  irregular  in  shape,  with  short, 
lobose  pseudopodia;  slowly  ameboid.  Body  colorless  and  transparent. 
Distinct  differentiation  of  protoplasm  along  the  entire  surface,  ectoplasm 
forming  outer  layer.  Endoplasm  faintly  granular,  with  numerous  small 
fat  globules.    Monosporous  and  disporous. 

Spore:  Central  portion  greatly  enlarged;  ovoid,  with  very  long  tapering 
process  extending  out  from  each  end.  Sutural  line  perpendicular  to  the 
longest  diameter.    Polar  capsules  large  and  spherical.    Sporoplasm  finely 


319]  STUDIES  ON  MYXOSPORIDIA—KUDO  81 

granular,  chiefly  located  in  one  valve,  extending  into  the  other  only  a  short 
distance  beyond  the  capsule.  Dimensions:  breadth  80/*,  breadth  of 
enlarged  central  portion  IS^t,  sutural  diameter  7/*,  diameter  of  polar  cap- 
sules 4)Lt. 

Genus    MYXOPROTEUS  Doflein  emend.     Davis 

1898        Myxoproteus  Doflein  1898  :  287 

1917        Myxoproteus  Davis  1917  :  219 

The  characters  of  the  genus  are  described  on  page  56. 
Type  species:  Myxoproteus  ambiguus  (Thelohan)  Doflein. 

MYXOPROTEUS  AMBIGUUS  (Thelohan)  Doflein 
[Figs.  75  to  80] 

1895        Myxosoma  atnbiguum  Th61ohan  1895  :  344 

1898        Myxoproteus  ambiguus  Doflein  1898  :  287-288 

Habitat:  Urinary  bladder  of  Lophius  piscatorius;  Le  Croisic,  Rovigno, 
Napoli. 

Vegetative  form:  Polymorphous.  Color  milky  white.  Protoplasm  is 
filled  with  numerous  granules  and  fat  globules.  Pseudopodia,  short, 
pointed  lobose.  Plasmogamy  and  plasmotomy  take  place.  Many  small 
individuals  formed  apparently  by  plasmotomy,  often,  make  up  groups. 
Disporous,  polysporous? 

Spore:  Almost  pyramidal,  with  anterior  processes.  Two  very  large 
polar  capsules  at  the  anterior  end.  The  distance  between  these  capsules 
is  equal  to  or  greater  than  the  diameter  of  the  capsules.  Sporoplasm  with 
two  nuclei.  Dimensions:  length  25/i,  breadth  18  to  20/x,  diameter  of  polar 
capsules  7/i. 

MYXOPROTEUS  CORDIFORMIS  Davis 

[Figs.  81  to  83] 

1917        Myxoproteus  cordiformis  Davis  1917  :  231 

Habitat:  Urinary  bladder  of  Chaetodipterus  faber;  Beaufort  (June, 
July). 

Vegetative  form:  Rounded;  very  slowly  ameboid,  usually  forming  a 
single,  short,  lobose  pseudopodium.  Body  colorless  and  transparent. 
Ectoplasm  not  distinct.  Entire  trophozoite  finely  granular,  with  a  few 
fat  globules.  Rarely  vacuolar.  Small  trophozoites  often  show  a  single 
large,  central  vacuole.  Rounded  sporulating  trophozoites  18/t  in  diameter. 
Disporous. 


82  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [320 

Spore:  Heart-shaped  in  front  view,  with  peculiar  wing-like  expansions 
on  each  side  which  contain  remains  of  parietal  cells.  Sutural  plane  oblique 
in  position.  Capsulogenous  cells  distinct.  Sporoplasm  finely  granular, 
fills  the  extracapsular  cavity  of  the  spore.  Dimensions:  length  \2n, 
breadth  10  to  ll/x,  thickness  6/1,  polar  capsules  3  to  4/i  in  diameter. 

MYXOPROTEUS  CORNUTUS  Davis 
[Fig.  84] 

1917        Myxoproteus  corntdus  Davis  1917  :  231 

Habitat:  Urinary  bladder  of  Bairdiella  chrysura;  Beaufort. 

Vegetative  form:  Somewhat  elongated  or  irregular  in  shape,  with 
short  lobose  pseudopodia;  slowly  ameboid.  Differentiation  of  protoplasm 
clear.  Ectoplasm  well  developed,  hyaline;  in  rounded  individuals  forming 
a  distinct  layer  around  the  body.  Endoplasm  opaque,  contains  coarse 
refringent  granules  varying  in  shape,  and  a  few  fat  globules.  In  contracted 
rounded  resting  condition,  endoplasm  becomes  condensed,  while  ectoplasm 
appears  more  abundant.  Rounded  trophozoites  up  to  27^  in  diameter. 
Disporous. 

Spore:  Heart-shaped,  with  two  anterior  processes.  Shell  very  thick. 
Polar  capsules  large,  opening  some  distance  apart.  Coiled  polar  filament 
distinct.  Sporoplasm  finely  granular,  with  a  few  small  fat  globules,  fills 
the  extracapsular  cavity  of  the  spore.  Dimensions:  sutural  diameter 
exclusive  of  the  processes  9/u,  breadth  12)u,  length  of  processes  5n,  diameter 
of  polar  capsules  3^1. 

Genus    WARDIA    nov.  gen. 

The  characters  of  the  genus  are  described  on  page  56. 
Type  species:  Wardia  ovinocua  nov.  spec. 

WARDIA  OVINOCUA  nov.  spec. 

[Figs.  85  to  95] 

Habitat:  Ovum  and  connective  tissue  of  ovary  of  Lepomis  humilis 
Girard;*  Salt  Fork,  111.  (November).  Only  one  fish,  6.5  cm.  long  with 
normal  appearance,  was  found  to  be  infected. 

Vegetative  form:  Trophozoites  form  cysts  visible  to  the  naked  eyes 
as  white  spherical  spots  in  the  pink-colored  ovary.  Four  cysts  present. 
The  cyst  (Figs.  85  and  86),  in  section,  shows  a  circular  form  surrounded  by 
several  layers  of  hypertrophied  nurse  cells  and  connective  tissue,  in  which 
many  large  blood  vessels  are  present.     Protoplasm  is  not  clearly  differ- 

*Professor  F.  Smith  of  the  Department  kindly  identified  all  the  fish  that  were  collected 
in  the  vicinity  of  Urbana  and  mentioned  in  this  paper  as  hosts,  for  which  the  writer  wishes  to 
express  his  appreciation. 


321]  ■  STUDIES  ON  MYXOSPORIDIA— KUDO  83 

entiated  into  ectoplasm  and  endoplasm,  the  whole  protoplasm  showing 
granulated  reticular  structure.  Cysts  contained  numerous  fully  developed 
spores  and  a  small  number  of  spores  in  developmental  stages,  which  sug- 
gested the  fact  that  two  spores  rise  from  each  pansporoblast.  The  parasite 
is  also  found  in  the  state  of  diffuse  infiltration  in  the  connective  tissue 
around  the  cyst.  Diameter  of  cysts  316  to  445/i  in  sections.  Polysporous. 
Spore:  In  front  view,  isosceles  triangular  form,  two  sides  of  which 
usually  convex,  with  more  or  less  attenuated  anterior  end  (Figs.  87,  90, 
92);  in  profile,  ellipsoidal  (Fig.  88);  and  oval  viewed  from  the  anterior 
end  (Fig.  89).  Sutural  plane  at  right  angles  to  the  longest  diameter  (Figs. 
87  and  89),  Shell  comparatively  thin  except  at  the  anterior  end  and  has 
many  fine  network-like  ridges  on  the  surface.  These  ridges  are  hardly 
observable  on  fresh  spores  on  account  of  their  fine  form  and  the  conspicu- 
ously large  polar  capsules  lying  in  the  spore.  When  stained,  however,  they 
are  not  only  made  distinctly  visible,  but  the  prolongation  of  each  ridge 
from  the  posterior  edge  wbiph  forms  about  l/x  long  fringe-like  structure  is 
also  more  clearly  recognized  (Figs.  90-95).  Two  large  and  spherical  polar 
capsules  located  in  the  central  portion  of  the  spore.  Coiled  (5  to  6  times) 
polar  filament  extremely  distinct.  The  openings  of  polar  capsules  at  the 
anterior  end.  Sporoplasm  comparatively  small,  finely  granular,  without 
any  vacuole,  contains  two  small  nuclei,  when  stained.  Dimensions  in  vivo: 
sutural  diameter  9  to  lO/x,  breadth  10  to  12/i,  thickness  6ju,  diameter  of  the 
polar  capsule  4/i,  length  of  polar  filament  35  to  45/x. 

WARDIA  OHLMACHERI  (Gurley)  Kudo 

[Figs.  96  and  97] 

1893        Myxosporidian  Ohlmacher  1893  :  561-567 

1893  Chloromyxum  ohlmacheri  Whinery  1893  :  660-662 

1894  Chloromyxum  {Spkaerospora) 

ohlmacheri  Gurley  1894  :  267-272 

1895  ?  Leptotheca  ranae  Th61ohan  1895  :  383 
1899        Leptotheca  ohlmacheri                 Labh€  1899  :  87 

Habitat:  Urinary  tubules  of  kidney  of  Bufo  lentiginosus  Shaw  and 
kidney  of  Rana  esculenta  and  R.  temporaria  {R.  fusca) ;  Sycamore,  De  Kalb 
county.  111. 

Vegetative  form:  Not  found. 

Spore:  Transversely  elliptic.  Sutural  plane  perpendicular  to  the  longer 
axis  of  the  spore.  A  well  defined  undulate-parallel  longitudinal  striation 
on  the  shell.  Sutural  ridge  comparatively  well  marked.  Two  polar  capsules 
lying  side  by  side,  occasionally  only  one.  Dimensions:  sutural  diameter 
6ju,  breadth  8/i,  diameter  of  polar  capsule  3  to  3.5/:,  length  of  polar  filament 
6  to  8  times  the  breadth  of  spore  (48  to  64/i). 

Remarks:  This  form  is  apparently  very  much  different  from  any 
species  of  genus  Leptotheca,  in  the  general  form,  form  of  polar  capsules, 


84  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [322 

striations  on  the  shell  and  the  habitat.  Tho  the  form  of  the  spore  is  differ- 
ent from  the  type  species  of  the  genus  Wardia  and  nothing  is  known  about 
the  vegetative  form,  the  presence  of  large  spherical  polar  capsules  in  the 
central  portion  of  the  spore,  the  striations  on  the  shell  and  the  occurrence 
of  the  same  nature,  i.e.,  from  fresh  waters  in  the  close-by  localities,  show 
its  nearer  relationship  to  the  genus  Wardia  than  to  the  genus  Leptotheca. 
Hence  it  is  placed  here  provisionally. 

Genus     MITRASPORA  Fujita     emend.  Kudo 
1912        Mitraspora  Fujita  1912  :  259-260 

The  characters  of  the  genus  are  described  on  page  56. 
Type  species:  Mitraspora  cyprini  Fujita. 

MITRASPORA  CYPRINI  Fujita 

[Figs.  98  to  104] 

1912        Mitraspora  cyprini  Fujita  1912  :  259-260 

Habitat:  Renal  tubules  of  kidney  and  ureters  of  Cyprinus  carpio  L. 
and  Carassius  auratus  L. ;  Sapporo  (winter),  Tokio  (March). 

Vegetative  form:  Fujita's  only  description  is  as  follows:  "The  sporo- 
blast  contains  generally  three  or  four  spores."  The  present  writer  observed 
a  similar  form  in  the  ureter  and  the  renal  tubule  of  the  kidney  of  Cyprinus 
carpio  L.,  in  Tokio,  The  observations  are  as  follows:  Trophozoites  small 
ameboid  (Fig.  98).  Body  colorless.  Movements  tardy.  Differentiation 
of  protoplasm  imperfect.  The  hyaline  ectoplasm  recognizable  at  one  side 
of  the  body,  where  lobose  pseudopodia  are  formed  (Figs.  98-99).  Endo- 
plasm  granular  with  vacuoles  and  brownish  granules,  which  become 
larger  as  the  body  grows.  Size  10  to  40/i.  Disporous  (Kudo)  and  poly- 
sporous  (?,  Fujita). 

Spore:  Fujita's  descriptions  are  as  follows:  Form  resembles  monk's 
hood,  slightly  attenuated  at  its  anterior  end.  Shell  uniformly  thin,  except 
at  two  points  of  the  truncated  posterior  end.  Each  shell  valve  has  eight 
distinct  striations  which  run  longitudinally  and  turn  into  long  cilia  up  to 
5.8jLi  long,  planted  in  a  single  row  at  the  posterior  end  of  the  spore.  Two 
polar  capsules  at  the  anterior  end.  The  nucleus  is  obscure  and  no  vacuole 
is  present.  Dimensions:  length  10  to  13/x,  breadth  Sn,  polar  capsules 
3.8/x  by  2n,  length  of  polar  filament  15/i  (weak  glycerine).  The  writer 
observed  the  following  facts:  More  rounded  with  rounded  anterior  end  in 
front  and  side  views.  Shell  more  or  less  thick  along  the  entire  posterior 
margin.  Striations  on  shell,  variable  in  number.  Sporoplasm  granular, 
without  any  vacuole,  exhibits  two  nuclei  when  stained.  Posterior  filaments 
5  to  6  in  number  and  5  to  6/i  long,  being  absent  in  some  spores.  Dimen- 
sions in  vivo:  length  lO/x,  breadth  8  to  9/i,  thickness  6  to  S/x,  polar  capsule 
4n  by  1.5  to  2/i,  length  of  polar  filament  35  to  40/i. 


323]  STUDIES  ON  MYXOSPORIDIA—KUDO  85 

Remarks:  Tho  Fujita  does  not  describe  the  vegetative  form  and  there 
are  some  differences  in  the  form  and  size  of  the  spore  between  the  forms, 
the  writer  does  not  find  out  any  objection  against  the  union  of  the  above 
mentioned  two  forms. 

f 

MITRASPORA  CAUDATA  (Parisi)     Kudo 

[Figs.  105  to  107] 


1910 

Sphaerospora  caudata 

Parisi 

1910 

:  253-254 

1912 

Sphaerospora  caudata 

Parisi 

1912 

:289 

1913 

Sphaerospora  caudata 

Parisi 

1913 

:  396-402 

Habitat:  Renal  tubules  of  kidney  of  Alosa  finta  Cuv.  var.  lacustris 
Fatio;  Lake  Como. 

Vegetative  form:  Rounded  or  variously  elongated  owing  to  the  move- 
ments. Protoplasm  is  distinctly  differentiated  into  ectoplasm  and  endo- 
plasm.  Ectoplasm,  hyaline  and  homogeneous,  forms  slowly  moving  lobose 
pseudopodia.  Endoplasm  granular,  contains  yellow  globules  and  fat 
granules.     Disporous  and  polysporous. 

Spore:  Subspherical  in  front  view;  oval  in  profile;  anterior  end  being 
more  rounded  than  the  posterior  end.  Shell  rather  thick,  longitudinally 
striated.  In  front  view,  the  posterior  end  enlarged  into  a  quadrangular 
form,  which  appears  as  a  small  spine  in  side-view  and  which  projects 
backward  long  and  fine  filaments,  usually  six  in  number.  Two  well  devel- 
oped polar  capsules  open  on  each  side  of  the  sutural  plane.  Polar  filament 
coiled  5  to  6  times.  Sporoplasm  without  any  iodinophilous  vacuole. 
Dimensions:  external  length  10  to  llju,  internal  length  7  to  9n,  length  of 
polar  capsules  4  to  4.5)u,  length  of  polar  filament  up  to  48/x,  length  of  pos- 
terior filaments  up  to  28/i. 

MITRASPORA  ELONGATA  nov.  spec. 
[Figs.  602  to  621] 

Habitat:  In  the  urinary  tubules  and  tissue  of  kidney  of  Lepomis 
cyanellus;  Crystal  Lake,  Urbana,  111.  From  June  to  July,  all  the  fish 
examined,  36  in  number  and  10  cm.  in  average  length,  were  found  to  be 
infected.  Other  fish  such  as  Lepomis  pallidus  and  Lepomis  humilis,  caught 
at  the  same  time,  were  free  from  the  infection.  Early  in  June,  the  number 
and  size  of  the  parasites  in  a  host  body  were  rather  small  and  only  a  small 
number  of  spores  could  be  recognized  in  the  fresh  state  with  the  addition 
of  potassium  hydrate  solution.  The  growth  of  the  parasite  was  rather 
remarkable  during  the  hot  weeks  in  the  latter  part  of  June  and  July  so 
that  every  fish  caught  on  July  17th  showed  a  heavy  infection,  exhibiting 
small  whitish  pustules  over  the  surface  of  the  organ.  During  June, 
vegetative  forms  and  spores  were  found  in  the  lumen  of  the  urinary  tubules. 


86  -  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [324 

altho  some  contained  the  parasitic  masses  in  the  tissue.  About  the  middle 
of  July,  the  parasite  forms  conspicuous  cysts  in  the  tissue  thruout  the 
organ.  The  c)^t  may  or  may  not  be  covered  by  a  thick  layer  of  connective 
tissue  from  the  host.  Aside  from  this  hypertrophy,  the  host  did  not  show 
any  pathological  change  which  covdd  be  recognized.  » 

Vegetative  form:  Youngest  trophozoite  found  in  the  urinary  tubule 
is  multinucleate,  rounded,  and  of  from  20  to  50ai  in  diameter.  The  proto- 
plasm is  not  differentiated,  the  entire  body  is  finely  granular  or  coarsely 
reticular  in  structure.  In  the  protoplasm  are  to  be  seen  nuclei  and  sporo- 
blasts  at  different  stages  of  development.  The  union  of  two  propagative 
ceUs  similar  to  that  of  Myxoholus  toyamai  produces  a  small  body  which 
developes  into  a  single  sporoblast  and  ultimately  into  a  single  spore 
(Figs.  605-613).  In  later  stages,  the  trophozoite  reaches  a  size  of  200/1 
in  diameter  showing  many  stages  of  spore  formation  and  mature  spores, 
surrounded  by  thick  layers  of  connective  tissue  from  the  host.  Poly- 
sporous. 

Spore:  Elongated  oblong  with  pointed  anterior  and  truncated  posterior 
extremities.  The  width  is  often  greatest  at  the  middle  of  the  polar  cap- 
sules, the  posterior  portion  is  much  narrower  than  the  anterior.  Nearly 
circular  in  the  cross-section  thru  the  polar  capsules.  The  shell  is  thin, 
the  sutural  line  being  faintly  marked  in  fresh  state.  It  generally  is 
obliquely  located  in  relation  to  the  capsules.  The  shell  also  shows  fine 
longitudinal  striations,  14  to  16  in  number,  on  each  valve.  The  sutural 
line  as  well  as  the  striations  are  best  seen  in  spores  stained  with  Heiden- 
hain's  iron  hematoxylin.  Two  polar  capsules  elongated  pyriform,  mostly 
equal  in  size,  occupy  the  anterior  half  of  the  spore.  Abnormal  situations 
of  the  polar  capsules  are  sometimes  observed  (Fig.  619).  The  coiled  polar 
filament  is  faintly  visible  in  fresh  spores.  It  is  spirally  coiled  along  the 
wall  of  the  polar  capsule  without  any  central  axis.  This  fact  was  clearly 
observed  in  stained  section  as  is  shown  in  Figs.  620  and  621.  The  fila- 
ment has  seven  or  eight  windings,  thus  agreeing  with  the  actual  length 
of  the  extruded  polar  filament.  The  polar  filament  was  extruded  under 
the  action  of  potassium  hydrate  solution.  The  extrusion  takes  place 
even  in  some  spores  which  were  treated  with  Schaudinn's  fixative  and 
kept  in  95  per  cent  alcohol  for  three  months  (see  the  similar  observations 
on  Myxoholus  discrepans  on  page  157).  The  sporoplasm  is  finely  granular 
and  transparent.  When  stained,  it  shows  two  nuclei  in  the  center  or  near 
the  posterior  part  of  the  body.  Dimensions  of  preserved  spores:  length 
15  to  17/ii,  breadth  5  to  6/i,  thickness  4.5  to  5.5/z,  polar  capsule  7.5^  by  2/x, 
length  of  polar  filament  40  to  50/Lt. 

Suborder    SPHAEROSPOREA     nom.  nov. 
The  definition  of  the  suborder  is  recorded  on  page  57. 


325]  STUDIES  ON  MYXOSPORIDIA—KUDO 

Family     CHLOROMYXIDAE     Thelohan 

1892        Chloromyxidees  Thfilohan  1892  :  173 

1895        ChloromyxidSes  Thaohan  1895  :  344 

The  characters  of  the  family  are  described  on  page  57. 

Genus     CHLOROMYXUM     Mingazzini 

1890        Chloromyxum  Mingazzini  1890  :  160 

1892        Chloromyxum  Thdlohan  1892  :  173-176 

1895        Chloromyxum  Thflohan  1895  :  344 

The  characters  of  the  genus  are  described  on  page  57. 
Type  species:  Chloromyxum  leydigi  Mingazzini. 

CHLOROMYXUM  LEYDIGI  Mingazzini 
[Figs.  108  to  113] 


87 


1851 

Leydig 

1851 

:  225-234 

1852 

Leuckart 

1852 

:435 

1854 

Lieberkiihn 

1854 

:352 

1890 

Chloromyxum  leydigi 

Mingazzini 

1890  : 

:  160-164 

1892 

Chloromyxum  leydigi 

Thelohan 

1892  ; 

:  166,  169-170 

1894 

Chloromyxum  leydigi 

Chloromyxum  incisum 

Gurley 

1894; 

:  25^260 

1895 

Chloromyxum  leydigi 

Chloromyxum  incisum 

Thelohan 

1895  : 

:  345-346 

1898 

Chloromyxum  leydigi 

Doflein 

1898  : 

:  292,  310,  etc. 

1912 

Chloromyxum  leydigi 

Erdmann 

1912 

:  149-162 

1916 

Chloromyxum  leydigi 

Georg6vitch 

1916a 

:3 

1917 

Chloromyxum  leydigi 

Davis 

1917 

:  236-237 

1917 

Chloromyxum  leydigi 

Erdmann 

1917 

:  276-321 

1918 

Chloromyxum  leydigi 

Georgfivitch 

1918 

:  182-189 

Habitat:  Gall-bladder  of  Rhina  squatina  L.,  Spinax  spinax  L.,  ScylUum 
canicula, S.asterias,Raja batis L.,  i?.  clavata L.,  R.  undulata Lac,  Torpedo narce 
Ris.,  T.marmorata,  T.ocellata,  T. torpedo  'L.,Acanthias  acanthias  L,.,Trygon 
pastinaca  L.,  Dasybatis  hastatus,  D.  sabina,  Pteroplatea  machira  Le  Sueur, 
Scoliodon  terrae-novae,  Cestracion  zygaena,  C.  tiburo,  Carcharhinus  limbatus; 
Roscoflf,  Concarneau,  Marseille,  Banyuls,  Rovigno,  Heligoland,  Beaufort, 
Monaco  (May),  Napoli,  Genova.  Erdmann  observed  the  species  at 
Naples  from  March  to  August.  She  noticed  mixed  infection  with  Cerato- 
myxa  reticulata  and  especially  with  Leptotheca  parva.  Georgevitch  studied 
the  parasite  at  Monaco  from  February  to  April. 

Vegetative  form:  Polymorphous,  being  spherical,  oval  or  irregular. 
The  change  of  the  form  rather  rapid  under  favourable  conditions.  Differ- 
entiation of  protoplasm  distinct.  Ectoplasm  with  pseudopodia  of  various 
form,  i.e.,  lobose,  filiform  or  intermediary;  short,  pointed  or  branched. 
Endoplasm  alveolar,  filled  with  yellowish  granules.  Doflein  observed  the 
plasmotomic  multiplication  of  young  trophozoites.  Polysporous.  Erd- 
mann's  observations  (1917)  may  be  summarized  as  follows:  Ameboid. 


88  II.UNOJS  BIOLOGICAL  MONOGRAPHS  [326 

Color  of  the  body  greenish  to  dark  green.  The  protoplasm  is  clearly  dif- 
ferentiated into  ectoplasm  and  endoplasm.  The  ectoplasm  is  hyaline  and 
covers  the  entire  surface  of  the  body.  It  appears  as  a  fine  fibrous  structure 
when  fixed  with  Bouin's  solution.  The  endoplasm  contains  besides  nuclei, 
two  kinds  of  spherules;  one  smaller  and  yellowish  "color-carriers"  and  the 
other  larger  and  light  to  dark  greenish  reserve  bodies.  The  color-carrier 
is  in  part  composed  of  myelin,  while  the  reserve  body  is  of  glycogenous 
nature.  The  infection  was  studied  experimentally  per  os:  young  tropho- 
zoites appeared  in  3  to  5  days  which  continued  to  10th  day,  various  tropho- 
zoites were  seen  in  13  to  19  da}^,  and  sporulating  individuals  were  first 
recognized  in  39  days  after  the  infection.  The  trophozoite  multiplies  in 
number  either  by  fission  or  by  budding.  It  usually  contains  enclosures 
which  seem  to  be  degenerating  erythrocytes.    Mictosporous. 

Spore:  Ovoidal.  Shell- valves  show  wide  edge  at  sutural  plane,  which 
is  attenuated  at  the  anterior  end  and  forms  a  quadrilateral  process  at  the 
posterior  end,  from  which  a  row  of  ciUa  grows.  Shell-valves  have  ridges 
(6  to  7),  which  run  parallel  to  the  posterior  margin.  The  striations  may 
vary  considerably.  Four  polar  capsules  at  the  anterior  end.  Dimensions: 
length  8/i.  Erdmann  gave  the  following  dimensions:  Spores  from  Torpedo 
marmorata  and  T.  ocellata:  length  6  to  9yL,  breadth  5/i,  polar  capsule  3^ 
by  2/i.  Those  from  Scyllium  asterias:  length  8  to  9n,  breadth  6/x,  polar 
capsule  2m  by  1/u.  Those  from  Raja  hatis:  length  7  to  8//,  breadth  5m, 
polar  capsule  2n  by  1^.  Length  of  polar  filament  20  to  30m  (absolute  alco- 
hol warmed  up  to  40**  C). 

CHLOROMYXUM  CAUDATUM  Thelohan 
[Fig.  114] 
1895        Chhromyxum  caudatum  Th61ohan  1895  :  346 

Habitat:  Gall-bladder  of  Molge  cristata  Laur.;  Vicinity  of  Rennes. 

Vegetative  form:  Body  yellowish  with  lobose  pseudopodia.  Proto- 
plasm finely  granular. 

Spore:  Oval  or  spheroidal.  Shell  enlarged  at  the  anterior  part,  having 
a  simple  or  bifurcated  tail-like  process,  as  in  Henneguya,  at  the  posterior 
end.  Dimensions:  total  length  18m,  length  8m,  breadth  6  to  7m,  length  of 
tail  10m. 

CHLOROMYXUM  QUADRATUM  Thelohan 


[Figs.  115  to  117] 

1891 

Keiffer 

1891 

111 

1893 

Pfeiffer 

1893 

81 

1895 

Chlorotnyxum  quadratum 

Thaohan 

1895 

347 

1912 

CMoromyxum  quadratum 

Parisi 

1912 

289 

1913 

Chloromyxum  quadratum 

Awerinzew 

1913a 

:155 

1913 

Chhromyxum  quadratum 

Fennor 

1913 

199 

327]  STUDIES  ON  MYXOSPORJDIA—KUDO  89 

Habitat:  Muscle  of  Syngnathus  acus  L.,  Trachurus  trachurus  L., 
Nerophis  aequorens  L.,  Callionymus  lyra  L.,  Coris  julis  L.,  Ariodes  polysta- 
phylodon,  kidney  of  Blennius  gattorugine  Brunn;  Helder,  RoscoflF,  Concar- 
neau,  Marseille,  Beira  (Africa),  Napoli  (summer). 

Vegetative  form:  Not  described  by  any  of  these  authors. 

Spore :  Quadrangular  pyramid  with  curved  edges  and  roundish  angles. 
Four  polar  capsules  at  the  anterior  end.  Dimensions:  length  6)u,  breadth 
S/i,  length  of  polar  filament  8  to  10^. 


CHLOROMYXUM  FLUVIATILE  Thelohan 
[Fig.  118] 

1892        Chloromyxum  fluviatile  Thdlohan  1892  :  173-176 

1895        Chloromyxum  fluviatile  Th61ohan  1895  :  346 

Habitat:  Gall-bladder  of  Leuciscus  cephalus  L.;  Paris. 

Vegetative  form:  Young  trophozoites  colorless;  adults  yellowish. 
Form  highly  variable.  Active  change  of  the  form  of  body.  Clear  diflFer- 
entiation  between  ectoplasm  and  endoplasm.  Ectoplasm  usually  recog- 
nizable at  one  end  of  the  body  where  lobose  pseudopodia  are  formed. 
Size  reaches  25  to  30^.    Polysporous. 

Spore:  Spherical,  generally  small.  Sutural  ridge  fairly  well  marked. 
Dimensions:  7  to  8/i  in  diameter. 


CHLOROMYXUM  MUCRONATUM  Gurley 

[Figs.  119  to  122] 

1854 
1879 
1882 
1883     ■ 

1893  Chloromyxum  mucronatum 

1894  Chloromyxum  mucronatum 

1908  Chloromyxum  mucronatum 

1909  Chloromyxum  mucronatum 

Habitat:  Urinary-bladder  and  kidney  of  Lota  lota  L.;  Bodensee,  other 
localities  not  mentioned. 

Vegetative  form:  Spherical,  elliptical  or  irregular.  Size  up  to  75/ii 
in  diameter.    Protoplasm  containing  irregularily  scattered  fat-like  globules. 

Spore:  Sharp-contoured;  subglobular,  mucronate  anteriorly.  Dimen- 
sions: length  8/z. 


Lieberkiihn 

1854; 

: 352-353 

Leuckart 

1879  ; 

:248 

Butschli 

1882  ; 

:  PI.  38  :  17 

Balbiani 

1883  ; 

; 201,  203 

Gurley 

1893  : 

:419 

Gurley 

1894; 

:  264,  265 

Auerbach 

1908  ; 

:456 

Auerbach 

1909a 

:71 

Balbiani 

1866  :  600-602 

Balbiani 

1867  :  275,276,  335 

Butsrhli 

1882  :  590 

Keiffer 

1890  :  559 

Henn^iuy  et  Th€Iohan 

1892  :  587 

Gurley 

1893  :  411 

Gurley 

1894  :  281 

Thaohan 

1895  :  347 

90  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [328 

CHLOROMYXUM  DIPLOXYS  (Gurley)  Thelohan 
[Figs.  123  to  125] 

1866 
1867 
1882 
1890 
1892 

1893  Cystodiscus  ?  diploxys 

1894  Cystodiscus  ?  diploxys 

1895  Chloromyxum  diploxys 

Habitat:  Abdominal  cavity  of  Tortrix  viridana  L.  (imago); 

Vegetative  form:  Trophozoites  form  spherical  cysts,  230  to  400ju 
in  diameter.  Cyst  membrane  rather  thick.  Protoplasm  containing 
brownish  granules,  and  fat-like  globules  (red  with  iodine). 

Spore:  Elliptic  or  slightly  flattened.  Sutural  line  straight,  forming  a 
ridge.    Two  polar  capsules  at  each  end. 

CHLOROMYXUM  PROTEI  Joseph 

1905  Chloromyxum  protei  Joseph  1905  :  450-451 
1907        Chloromyxum  protei                   Joseph  1907  :  398-412 

Habitat:  Renal  tubules  of  kidney  of  Proteus  anguineus  L.;  Vienna. 

Vegetative  form:  Generally  rounded  or  sausage  form.  No  clear 
differentiation  between  ectoplasm  and  endoplasm.  Movements  slow. 
Probable  occurrence  of  plasmotomy  by  budding  and  division.  Size: 
40  to  45m  by  28  to  40m. 

Spore:  Spherical.  Shell  finely  striated  parallel  to  the  sutural  line. 
Four  polar  capsules  each  with  an  independent  opening.  Dimensions: 
10  to  13m  in  diameter,  polar  capsules  4  to  6m  long.  The  polar  filament 
appears  to  be  rather  short. 

CHLOROMYXUM  TRUTTAE  Leger 

[Fig.126] 

1906  Chloromyxum  truUae  L6ger  1906  :  267-270 
Habitat:  Gall-bladder  and  gall-duct  of  Truttafario  L.;  Dauphine. 
Vegetative  form:  Ameboid  form.     Elongated.     Form  resembles   an 

Amoeba  Umax  of  about  40m  in  length.  Roundish  or  irregularly  contoured, 
with  small  pseudopodia.  Ovoidal  or  spherical,  25  to  40m  in  diameter 
without  any  visible  pseudopodia  (resting  state).  Body  colorless,  clear  and 
hyaline.  Very  active  movements  which  last  for  several  hours  after  the 
death  of  the  host.  Broad  and  obtuse  pseudopodia  well  developed  at  the 
anterior  end  of  the  body.  Endoplasm  alveolar,  contains  variable  numbers 
of  nuclei,  which  are  seen  in  vivo,  refractive  bodies  and  chromatic  granules. 
Monosporous(?)  and  polysporous. 

Spore:  Spherical.  Four  polar  capsules  of  different  size.  Shell- valves 
marked  with  parallel  ridges.    Dimensions:  8  to  9m  in  diameter. 


329]  STUDIES  ON  MYXOSPORIDIA—KUDO  91 

CHLOROMYXUM  CRISTATUM  Leger 

[Figs.  127  and  128] 

1906        Chloromyxutn  crishUum  L^ger  1906  :  270-272 

Habitat:  Gall-bladder  of  Tinea  vulgaris  Cuv.;  Grenoble. 

Vegetative  form:  Ordinarily  massive,  with  oval  or  round  contours, 
without  noticeable  pseudopodium.  Ectoplasm  hyaline.  Endoplasm 
granular  and  colorless.  Average  diameter  of  the  adults  about  20^.  Mono- 
sporous,  rarely  disporous.- 

Spore:  Spherical  or  subspherical.  Ten  marked  ridges  run  antero- 
posteriorly  on  each  shell- valve,  so  that  it  presents  a  cog-wheel  form  in  cross 
section.  Four  polar  capsules  at  the  anterior  end,  one  pair  being  smaller 
than  the  other.    Sporoplasm  with  two  nuclei.    Dimension:  10  to  ll/u. 

CHLOROMYXUM  DUBIUM  Auerbach 
[Figs.  129  to  133] 
1908        Chloromyxutn  dubium  Auerbach  1908  :  456-459 

1910        Chloromyxutn  dubium  Auerbach  1910c  :  177 

Habitat:  Gall-bladder  of  Lota  vulgaris  Cuv.;  Bodensee  (April  to  Sep- 
tember). 

Vegetative  form:  Spherical  or  rounded.  Rarely  irregular  forms. 
Protoplasm  is  dififerentiated  distinctly  into  ectoplasm  and  endoplasm. 
Ectoplasm  very  thin,  forms  pseudopodia  which  move  slowly.  Endoplasm 
granular,  contains  fat  globules.  Majority  of  the  trophozoites  appear  to 
live  floating  in  the  bile,  while  some  are  attached  to  the  epithelium  of  the 
bladder.    Disporous  and  polysporous. 

Spore:  Spherical,  with  four  polar  capsules.  Each  shell  valve  has 
longitudinal  ridges,  variable  in  number  (6  ridges  are  found  on  the  drawing), 
which  run  parallel  to  the  sutural  line.  Four  polar  capsules  of  nearly  same 
size  and  convergent.  Sporoplasm  finely  granular  with  two  nuclei.  Dimen- 
sions: diameter  10.8/i,  length  of  polar  capsule  3.6/i. 

CHLOROMYXUM  sp.  Awerinzew 

[Fig.  134] 

1908        Chloromyxum  sp.  Awerinzew  1908  :  43, 47, 48 

Habitat:  Gall-bladder  of  Raja  radiata;  Murman  coast?. 

Vegetative  form:  Form  rounded.  The  protoplasm  is  distinctly  differ- 
entiated into  ectoplasm  and  endoplasm.  Ectoplasm  hyaline  and  compara- 
tively abundant  in  quantity  compared  with  endoplasm,  forms  lobose 
pseudopodia  of  active  movements.  Endoplasm  vacuolated,  contains 
enclosures.  Between  the  two  layers,  a  thin  layer  of  protoplasm,  reticular  in 
structure  and  stained  deeply  with  hematoxylin,  is  present. 

Spore:  No  figure. 


92  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [330 

CHLOROMYXUM  THYMALLI  Lebzelter 

1912  Chloromyxum  thymalli  Lebzelter  1912  :  295-296 
Habitat:  Gall-bladder  of  Thymallus  thymallus  L.;  Vienna? 
Vegetative  form :  Irregular  form,  ?)2)  to  35ju  long  in  average.    Endoplasm 

contains  fat  globules  which  stain  brown  with  carmine.  Trophozoites 
attached  to  the  epithelium.  In  average,  6  spores  formed  in  each  individual. 
Intracellular  stage  in  the  epithelial  cell  is  supposed.    Polysporous. 

Spore:  Spherical.  Shell  structure  similar  to  C.  protei,  but  ridges  are 
more  developed  and  exhibit  somewhat  wavy  courses.  Polar  capsules  of 
equal  size.    Dimensions:  9  to  9.5ju  in  diameter,  polar  capsules  3^. 

CHLOROMYXUM  KOI  Fujita 
[Fig   135] 

1913  Chloromyxum  koi  Fujita  1913  :  257-259 

Habitat:  Gall-bladder  of  Cyprinus  carpio  L.;  Sapporo  (Nippon). 

Vegetative  form :  Spherical,  with  greatest  diameter  lip  to  SOju,  contain- 
ing 1  to  3  spores.  Each  spore  is  situated  in  a  clear  space  surrounded  by  a 
membranous  envelope  (sporoblast?),  around  which  there  is  some  finely 
granular  matter  (endoplasm?). 

Spore:  Spherical,  exhibiting  a  somewhat  angular  contour  at  the  ante- 
rior end.  Shell  thick  and  has  well  marked  ridges  on  the  surface,  i.e.,  4  to  5 
circular  ridges  and  on  both  sides  of  these  ridges,  two  more  ridges  each  bent 
in  a  loop-like  manner,  so  that  the  outline  of  spore  in  cross  section,  is  very 
much  like  of  a  toothed  wheel  with  nearly  equidistant  teeth,  16  to  18  in 
number.  Four  polar  capsules,  two  slightly  larger  than  the  other  two. 
Dimensions:  length  16)u,  breadth  lO/x,  length  of  polar  capsule  4/i,  length  of 
polar  filament  64/x. 

CHLOROMYXUM  MAGNUM  Awerinzew 

[Figs.  136  to  138] 

1913        Chloromyxum  magnum  Awerinzew  1913  :  155-156 

Habitat:  Gall-bladder  of  Acanthias  blainvillei;*  Algoa  Bay,  East  Lon- 
don, Liideritzbucht  (Africa). 

Vegetative  form:  Ameboid.  Body  yellowish  by  the  presence  of  large 
yellowish  granules  in  endoplasm.  Often  round  or  rosary  form.  Pseudo- 
podia  sometimes  absent,  so  that  the  trophozoites  move  like  Amoeba  Umax 
with  a  cluster  of  small,  hairy  pseudopodia  at  the  posterior  end.  In  larger 
form,  small  round  pseudopodia,  composed  of  homogeneous  ectoplasm,  are 
formed.  Plasmotomj'^  by  budding,  was  often  observed.  Usually  polyspor- 
ous, rarely  monosporous. 

Spore:  Elongated  spherical  form.  Four  polar  capsules  at  the  narrow, 
anterior  end.  Sporoplasm  with  two  nuclei.  Dimensions:  length  40  to 
48/1,  breadth  30  to  38/x,  length  of  polar  capsules  12  to  ISfi. 

*  Misprinted  in  Awerinzew's  paper  as  blainvilei. 


331]  STUDIES  ON  MYXOSPORJDIA—KUDO  93 

CHLOROMYXUM  FUNDULI  Hahn 
[Figs.  139  and  140] 

1915  Chloromyxum  funduli  Hahn  1915:205-206 

Habitat:  Muscle  of  Fundulus  sp.;  Woods  Hole.    In  one  fish. 

Vegetative  form:  Hahn  made  observations  on  few  fresh  and  stained 
smears.  According  to  him,  it  is  clear  that  the  staining  was  abnormal. 
It  is  hard  to  quote  this  here  as  he  used  different  terms  without  giving  any 
definition.    The  reader  is  advised  to  consult  Hahn's  paper. 

Spore:  Form  slightly  resembles  that  of  Choloromyxum  quadratum. 
Posterior  end  rounded,  the  anterior  portion  narrow  and  truncated  at  the 
tip;  optical  cross-section  thru  the  posterior  part  of  the  polar  capsules, 
circular.  Four  polar  capsules  at  the  anterior  end.  Dimensions:  height 
(length)  6)u,  breadth  and  thiclcness  7.5^  respectively. 

Remarlis:  As  to  the  comparison  of  the  present  species  with  Chloro- 
myxum clupeidae  Hahn,  see  p.  94. 

CHLOROMYXUM  MISGURNI  Kudo 
[Figs.  141  to  146] 

1916  Chloromyxum  tnisgurni  Kudo  1916  : 6-7 

Habitat:  Gall-bladder  of  Misgurnus  anguillicaudatus  Cantor;  Tokio 
(September). 

Vegetative  form:  Round  or  irregular.  Semicircular  when  viewed  from 
side.  From  the  flat  surface,  many  fine  root-like,  filiform  pseudopodia  are 
extruded.  No  clear  differentiation  between  ectoplasm  and  endoplasm. 
Endoplasm  alveolar.  Trophozoites  always  found  attached  to  the  lining 
epithelial  cells.  Size  up  to  50/x  by  20/i.  Polysporous  (6  to  8  spores),  rarely 
disporous. 

Spore:  Spherical,  slightly  attenuated  at  the  anterior  end.  Sutural  line 
straight  and  forms  a  ridge.  Fine  longitudinal  striations  run  parallel  to 
the  sutural  line.  Four  polar  capsules  at  the  anterior  end.  Sporoplasm 
finely  granular,  has  two  nuclei  of  equal  size.  Dimensions:  length  8  to 
9)u,  breadth  6  to  7/x,  thickness  5  to  6^,  length  of  polar  capsule  2  to  3ai, 
of  polar  filament  28  to  35m  (KOH). 

Remarks :  The  host  is  often  infected  at  the  same  time  by  Chloromyxum 
fujitai,  the  trophozoites  of  which  can  be  distinguished  from  the  present 
form  by  the  structure  and  the  floating  habit  in  the  bile.  Spores  in  the 
two  species  are  decidedly  different  in  form,  structure  and  size. 

CHLOROMYXUM  FUJITAI  Kudo 
[Figs.  147  to  152] 
1916        Chloromyxum  fujitai  Kudo  1916:7-9 

Habitat:  Gall-bladder  of  Misgurnus  anguillicaudatus  Cant.;  Tokio, 
(5%  of  the  fish  examined  in  September,  found  infected). 


94  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [332 

Vegetative  form:  Round  or  irregular.  No  clear  differentiation  of 
protoplasm.  Endoplasm  highly  vacuolated.  Ectoplasm  being  hardly 
distinguishable.  Size  up  to  40/t  in  diameter.  Trophozoites  float  in  the 
bile  in  almost  all  cases.    Disporous  and  polysporous  (up  to  8  spores). 

Spore:  Spherical,  often  attenuated  at  the  anterior  end.  Sutural  line 
not  straight.  Shell  very  thick,  shows  thick  ridges  running  longitudinally 
on  the  surface.  In  optical  cross  section,  the  spore  presents  an  outline  like 
a  cog-wheel  with  20  to  22  ridges.  The  thickness  of  ridges  varies  regularly; 
the  thickest  ones  being  located  on  two  lines  where  a  plane  perpendicular 
to  sutural  plane  cuts  the  shell  longitudinally,  others  decreasing  in  thick- 
ness as  they  approach  the  sutural  line.  Four  polar  capsules  at  the  anterior 
end.  Sporoplasm  with  two  nuclei.  Dimensions:  length  10  to  12jLt,  breadth 
8  to  10/i,  polar  capsules  2  to  3n,  length  of  polar  filament  23  to  SOn  (KOH). 

CHLOROMYXUM  CLUPEIDAE  Hahn 
[Figs.  153  to  156  and  562  to  565] 

1900  Sporozoa  Tyzzer  1900  :  66-^ 

1901  Sporozoa  Linton  1901  :  438 
1910  Chloromyxum  sp.  Auerbach  1910  :  178 
1917        Chloromyxum  dupeidae             Hahn  1917  :  15-19 

Habitat:  Body  musculature  of  Clupea  harengus,  Pomolobus  pseudohar- 
engus,  P.  aestivalis,  P.  mediocris,  Brevoortia  tyrannus,  Stenototnus  chrysops, 
Tautogolahrus  adspersus;  Woods  Hole. 

Tyzzer  mentioned  in  his  paper  and  also  in  a  letter  to  the  writer  that 
he  collected  the  material  in  August  of  1900  and  that  he  found  the  infection 
occurred  only  among  young  fish.  Hahn  also  called  attention  on  the  latter 
fact. 

Vegetative  form:  Hahn's  observations  are  as  follows: 

Clusters  of  spores  ("pseudocysts")  are  spindle-shaped,  especially  when 
young,  usually  l5ang  between  the  bundles  of  muscle  fibres.  Color  white  or 
creamy.  Larger  ones  usually  "in  pocket  just  beneath  the  integument." 
Schizogonic  multiplication  probably  exists.  Parasites  hard  to  stain, 
anilin  dyes  being  unable  to  stain  at  all.  Large  form  (probably  composed  of 
many  individuals)  890ju  by  30jli. 

Tyzzer  described  as  follows:  Cysts  up  tO"  1  to  2  mm.  in  length,  lying 
between  the  muscle  fibres  of  the  myotomes,  surrounded  at  times  by  mem- 
braneous connective  tissue.  The  parasites  also  occur  in  diffused  infiltration. 

Linton  found  two  cysts,  1.74mm.  by  1.16mm.  and  1.16mm.  by  0.58mm. 
and  also  diffused  state  between  the  fibrillae. 

The  writer's  observation  on  slides  prepared  by  Dr.  Tyzzer*  is  as  fol- 
lows:    Two  cysts  in  sections;  one  almost  spherical,  480/x  by  430/x,   sur- 

*The  writer  had  recently  the  opportunity  of  examining  the  slides  of  the  parasites  prepared 
by  Dr.  Tyzzer,  which  occasion  he  appreciated  very  much.  As  a  result  of  this,  the  writer 
became  convinced  of  the  identity  of  forms  observed  by  Tyzzer,  Linton  and  Hahn,  tho  he 
could  not  examine  the  latter  authors'  specimens. 


333]  STUDIES  ON  MYXOSPORJDIA—KUDO  95 

rounded  by  several  layers  (about  lO/i  thick)  of  connective  tissue  of  the 
host,  the  other  oval,  120^  by  110^.  The  staining  sufficed  to  reveal  only 
indistinct  structure  of  the  parasites.  The  homogeneous  ectoplasm  sur- 
rounds the  entire  surface  of  the  body  as  a  uniform,  but  very  thin  layer. 
Endoplasm  granular,  filled  with  spores  of  remarkably  identical  stages  of 
development.  Isolated  spores,  also,  occur  in  the  muscle  bundle  in  the 
state  of  diffused  infiltration.    Polysporous. 

Spore:  Hahn  describes  it  as  follows:  Low  conical  pyramid  with  round 
base;  square  with  bulging  sides.  No  indication  of  valves  in  the  spore 
shell.  Dimensions:  height  (length)  5)u,  breadth  and  thickness  7/*,  polar 
capsule  2/x  by  l^i. 

Linton's  form:  squarish  in  outline  with  rounded  corners,  7/i*  in  di- 
ameter. 

Tyzzer  describes  his  form  as  follows:  Quadrilateral  in  anterior  end 
view;  oval  in  side  view.  The  four  corners  are  a  little  protuberant  and  are 
directed  slightly  forward.  Shape  varies  considerably  in  different  species 
of  host.  The  corners  of  the  spore  from  Stenotomus  chrysops,  are  greatly 
drawn  out,  exhibiting  stellate  form.  Four  polar  capsules  radiating  from 
the  anterior  extremity  toward  the  four  corners.  Shell  shows  four  furrows 
radiating  from  the  anterior  extremity  outwards  to  the  side.  Sporoplasm 
occupies  extracapsular  cavity.  Polar  filaments  are  extruded  under  the 
action  of  acetic  acid.    Dimension:  breadth  7  to  7.5/t. 

The  writer's  observations  are  as  follows: 

Spores  in  fixed  and  decolorized  smears.  In  smear,  most  of  the  spores 
are  seen  lying  on  the  base  exposing  the  anterior  end  view  toward  the 
observer's  eyes,  a  few  lying  with  the  sutural  diameter  parallel  to  the  surface 
of  the  slide.  Form  quadrilateral  with  corners  more  or  less  drawn  out  in 
anterior  end  view;  oval,  with  concave  posterior  side  in  front  view  (Figs. 
562  to  564).  Shell  apparently  thin  but  was  not  clearly  separated  from  the 
sporoplasm  which  is  finely  granular  and  fills  the  extracapsular  cavity  of 
the  spore.  Four  polar  capsules  of  nearly  same  size  and  pyriform.  Coiled 
polar  filament  indistinct.  When  stained,  the  polar  capsules  stained  deeply. 
It  is  remarkable  to  see  almost  all  of  the  spores  exhibit  four  deeply  stained 
nuclei  of  capsulogenous  cells,  which  in  ordinary  case  disappear  as  the  spore 
matures.  Dimensions:  height  (length)  4  to  4.75)u,  breadth  and  thickness 
5.4  to  6.5ju,  polar  capsule  about  1.5/1  by  0.75)li. 

Remarks:  Thus  the  forms  of  Tyzzer,  Linton  and  Hahn  had  better 
be  treated  as  one  and  the  same  species.  As  to  the  distinction  of  Chloro- 
myxum  funduli  and  the  present  species,  the  writer  is  unable  to  make  it 
clear  as  he  could  not  examine  the  preparation  of  the  former  species  and 
especially  as  he  observed  some  intermediate  forms  between  these  two 
forms  in  Dr.  Tyzzer's  preparations  of  the  present  species. 


96  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [334 

CHLOROMYXUM  GRANULOSUM  Davis 

[Figs.  157  and  158] 
1917        CUoromyxum  granulosum         Davis  1917  :  237 

Habitat:  Urinary  bladder  of  Tylosurus  marianus;  Beaufort  (July, 
August). 

Vegetative  form:  Elongated  when  first  placed  on  the  slide,  but  soon 
becomes  contracted  and  motionless;  progressing  by  very  slow  ameboid 
movements.  Ectoplasm  usually  undistinguishable,  being  noticed  only  in 
a  few  individuals  which  had  formed  one  or  two  short,  lobose  pseudopodia 
of  hyaline  ectoplasm.  Body  colorless  to  light  yellow.  After  being  on  the 
slide  for  some  time  rounded  trophozoites  often  became  surrounded  by  a 
distinct  ectoplasmic  layer.  Entire  body  usually  coarsely  granular,  the 
granules  var\dng  greatly  in  size  and  shape;  sometimes  indistinctly  vac- 
uolated. Fat  globules  also  present.  Size  of  rounded  trophozoites  about 
30/i.    Disporous  and  polysporous. 

Spore:  Spherical,  with  four  distinct  ridges  on  the  posterior  half  of 
each  valve  converging  toward  the  anterior  end.  Sutural  ridge  distinct. 
Polar  capsules  pyriform  and  convergent.  Dimensions:  diameter  7/i, 
polar  capsules  2n. 

Remarks:  Trophozoites  from  some  fish  were"  all  colorless,  while  the 
larger  trophozoites  from  others  were  distinctly  yellow. 

CHLOROMYXUM  TRIJUGUM  nov.  spec. 
[Figs.  159  to  182] 

Habitat:  Gall-bladder  of  Lepomis  megalotis  Raf.;  Stony  Creek,  and 
Homer  Park,  HI.  (November).  The  parasite  was  only  found  in  this  species, 
Lepomis  humilis  and  L.  cyaneUus  seined  at  the  same  time  being  free  from 
the  infection.  Six  specimens,  three  from  each  of  the  above  mentioned 
locaUties,  harboured  abundantly  both  free  spores  and  trophozoites  of 
various  stages  of  development  in  the  bile.  The  fish,  from  6.5  to  10.5cm 
long,  were  normal  in  external  appearance  and  the  bladders  did  not  show  any 
particular  abnormality,  compared  with  those  of  other  fish,  as  is  usually 
the  case. 

Vegetative  form:  Trophozoites  float  usually  free  in  the  bile,  younger 
forms  are  most  frequently  attached  to  the  epithelium  of  the  bladder.  Form 
extremely  polymorphous,  manifesting  various  shapes  such  as,  almost  circu- 
lar, rounded,  oval,  elongated  or  irregular,  which  is  chiefly  due  to  the  active 
extrusion  and  retraction  of  the  pseudopodia  from  the  body  surface.  Body 
is  highly  transparent  and  colorless  in  both  the  young  and  the  adult.  The 
differentiation  of  protoplasm  into  ectoplasm  and  endoplasm,  is  distinctly 
visible  in  vivo  as  well  as  in  stained  preparations,  especially  in  larger  forms 
(Figs.  159  to  165).    The  endoplasm  presents  an  alveolar  structure  without 


335]  STUDIES  ON  MYXOSPORIDIA—KUDO  97 

any  enclosure  except  the  nuclei  and  various  stages  of  spore  formation 
(Figs.  161,  165,  168  to  171),  the  alveolar  network  being  smaller  at  the 
periphery  than  in  the  center.  The  ectoplasm  is  a  hyaline,  transparent  and 
homogeneous  layer,  free  from  any  course  granulation  in  fresh  conditions. 
It  shows,  however,  a  very  fine  reticular  structure  in  stained  preparations. 
The  pseudopodia  are  of  two  kinds  in  form,  always,  formed  of  ectoplasm 
alone :  the  filose  and  bristle-like  form,  sometimes  branching  and  protruding 
from  the  entire  surface  or  from  a  localized  part  of  the  body,  vary  in  length 
from  0.5  to  4/*  according  to  the  size  of  the  individual  (Figs.  159,  161,  164). 
This  form  developed,  sometimes,  into  a  thicker  form  with  two  to  four 
branched  finer  processes.  The  blunt,  lobose  pseudopodium  formed  at  a 
localized  part  of  the  body  is  well  recognizable  in  larger  individuals.  Fre- 
quently the  filose  and  the  lobose  pseudopodia  are  formed  on  a  trophozoite 
at  the  same  time.  The  movements  of  the  blunt  pseudopodia  were  striking 
in  some  specimens.  At  the  beginning  of  the  observation,  ten  minues  after 
the  bile  was  removed  from  the  host,  two  club-shaped  pseudopodia  (Figs. 
161  to  163)  which  were  extruded  from  a  trophozoite,  the  largest  diameter  of 
which  being  20/x,  moved  very  actively  in  the  semicircular  area  changing  their 
forms,  showing  maximum  length  of  20/x.  In  about  thirty  minutes,  they 
were  retracted  and  from  the  same  place,  a  short,  oval-shaped  pseudopodium 
was  seen  to  be  extruded,  which  remained  in  the  same  position  for  some 
time  without  great  change  of  form  (Fig.  164).  In  another  case,  a  tropho- 
zoite with  a  very  broad  and  rounded  pseudopodium  extruded  actively  two 
to  three  rounded  smaller  processes  at  its  extremity  (Figs.  165  to  167). 
After  fifteen  minutes  the  pseudopodium  was  retracted,  the  ectoplasm 
forming  a  uniformly  thick  layer  around  the  endoplasm.  The  observations 
were  done  at  room  temperature  in  hanging  drop  preparations,  sealed  with 
vaseline  and  paraffin,  by  using  comp.  oc.  12  and  apo.  imm.  ob.  2mm,, 
which  caused  no  mechanical  pressure  upon  the  parasites.  The  change  of 
form  and  especially  that  of  pseud«podia,  was  clearly  observed  for  one  hour 
and  twenty  minutes  under  the  above  mentioned  conditions  after  the  bile 
was  removed  from  the  host.  The  trophozoites  when  kept  for  sixteen  hours 
at  room  temperature,  underwent  degeneration  and  disintegrated,  setting 
free  the  spores  which  were  formed  in  them. 

No  active  multiplication  by  plasmotomy,  was  observed  in  vivo.  In 
fixed  preprations,  however,  forms  that  suggested  the  occurrence  of  the  pro- 
cess in  the  present  myxosporidian,  were  recognized.  As  was  stated  before, 
the  pseudopodia  are  always  formed  of  the  ectoplasm  and  as  each  portion 
of  these  dividing  forms  has  many  nuclei,  the  author  is  inclined  to  record 
the  presence  of  plasmotomy  in  the  present  form. 

Size  varies  greatly.  The  monosporous  form  10/x  by  14/i,  disporous 
15)Lt  by  25)Lt  and  polysporous  30)u  by  50/i,  the  largest  individual,  developing 
and  containing  more  than  200  spores,  was  300/x  by  50/i. 


98  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [336 

Spore:  Generally  circular  in  front  view;  oval  in  side  view.  Shell  com- 
paratively thick,  consequently  the  coiled  polar  filament  is  frequently  indis- 
tinct. Sutural  ridge  straight  and  distinct.  Each  valve  has  a  thick  straight, 
sometimes  slightly  zigzag-form  ridge  that  runs  parallel  to  the  sutural  line, 
so  that  in  side  view,  three  distinct  ridges  encircling  the  spore  are  recognized 
(Figs.  177  and  180),  From  each  of  these  two  ridges,  eight  to  twelve  short 
ridges  are  directed  toward  the  center  of  each  valve,  which  can  distinctly 
be  observed  on  the  spores  stained  with  Heidenhain's  iron  hematoxylin 
(Figs.  179  and  180).  They  can  be  seen  as  faint  markings  rising  from  the 
margin  directed  toward  the  center  of  the  spore,  in  front  view  of  fresh  spores. 
Four  pyriform  polar-  capsules  of  slightly  different  size  open  their  foramina 
independently  at  the  anterior  end  of  the  spore  (Figs.  178  and  181).  The 
sporoplasm,  granular  and  finely  reticular,  shows  almost  always  two  nuclei 
when  stained.  Dimensions  in  vivo:  length  and  breadth  8  to  10/i,  thickness 
5  to  In,  polar  capsules  3  to  Sn  by  2  to  3/x,  length  of  polar  filament  32  to 
40/i  (H2O2,  KOH). 

Remarks:  In  carefully  made  smear  of  the  bile,  a  number  of  empty 
spores  which  had  been  seen  in  fresh  hanging  drop  preparations,  and  often 
spores,  in  which  the  sporoplasm  with  two  elongated  nuclei  seemed  to  leave 
the  shell  (Fig.  182),  were  recognized.  As  this  particular  spore  was  found 
close  to  a  thicker  mass  of  the  wall  of  the  gall-bladder  in  the  smear,  it  can 
hardly  be  thought  that  the  mechanical  pressure  during  the  preparation 
lead  to  the  mission  of  the  sporoplasm  from  the  spore.  It  is  possible,  on  the 
other  hand,  to  think  that  this  is  one  of  the  cases  of  the  germination  of  the 
spore  in  the  host  in  which  they  were  developed,  as  was  reported  by  the 
author  in  Nosema  bombycis  Nageli  (Kudo,  1916). 

CHLOROMYXUM  CATOSTOMI  nov.  spec. 
[Figs.  560  and  561] 

Habitat:  Gall-bladder  of  Catostomus  commersonii  Lac;  Salt  Fork, 
Urbana,  111.  (October).    Four  fish,  from  8  to  14cm.;  apparently  normal. 

Vegetative  form:  Form  usually  rounded,  with  fiUform  pseudopodia. 
Majority  attached  to  the  epithelium,  a  few  being  free  in  the  bile.  Body 
colorless.  Protoplasm  is  not  well  differentiated.  Endoplasm  occupying 
the  entire  body  is  of  granular  structure  with  vacuoles  and  refringent 
spherules.  Size:  from  15  to  35)li.  When  kept  for  16  hours  in  a  refrigerator, 
the  trophozoites  liberated  the  spores.  The  number  of  spores  in  each  tro- 
phozoite is  usually  2  or  3,  rarely  5  to  6.  Active  plasmotomic  multiplication 
observed  when  examined.  Spores  were  comparatively  small  in  number, 
while  the  trophozoites  were  attached  abundantly  to  the  epithehum  of  the 
gall-bladder.    Disporous  and  polysporous. 

Spore:  Form  approximately  spherical  in  front  view;  oval  in  profile. 
Shell  with  very  fine  striations  which  run  parallel  to  the  sutural  ridge  that 
is  fairly  well  marked.    Rounded  polar  capsules  almost  of  same  size,  have 


3371  STUDIES  ON  MYXOSPORIDJA—KUDO  99 

independent  openings  at  the  anterior  end.  Coiled  polar  filament  indis- 
tinct. Abnormal  spores  with  five  polar  capsules  are  sometimes  seen. 
Dimensions  of  fixed  spores:  length  8j«,  breadth  7/i,  thickness  5  to  6fi,  polar 
capsules  2  to  2.5/x  by  l.S/i. 

CHLOROMYXUM  WARDI  nov.  spec. 
[Figs.  632  to  642] 

Habitat:  In  the  gall-bladder  of  Oncorhynchus  nerka:  Klutina  Lake, 
Alaska  (August).  A  single  gall-bladder  collected  and  preserved  in  formol 
by  Professor  Ward,  was  found  to  harbor  the  present  species.  The  study 
was  done  on  preserved  material  and  on  stained  smear  preparations. 

Vegetative  form:  Young  trophozoites  (Fig.  632)  show  ameboid  form, 
and  are  mostly  multinucleated.  The  protoplasm  is  not  well  differentiated 
either  in  unstained  material  or  in  stained  specimens.  It  is  granulated 
thruout  the  body,  and  is  vacuolated  at  places.  The  smallest  form  mea- 
sured was  18m  in  largest  diameter.  The  shape  of  the  body  suggests  its 
possession  of  ameboid  movements  when  alive,  altho  the  writer  could  not 
examine  fresh  specimens.  Large  trophozoites  in  which  the  spore  forma- 
tion had  partly  been  completed  are  generally  rounded  with  reticular  proto- 
plasm. Size  varies  to  some  extent.  The  trophozoite  shown  in  figure  633 
contains  six  mature  spores  and  is  23ai  in  largest  diameter.  The  largest 
one  found  was  38  by  30ju,  showing  ten  spores  and  nuclei.  Each  spore 
appears  to  develop  independently  from  a  single  sporoblast.  Disporous 
and  polysporous. 

Spore:  Rounded  pyramidal  in  front  view  (Figs.  640  and  641);  circular 
in  transverse  section  (Fig.  638).  The  shell  is  thickened  near  the  posterior 
margin  (Figs.  640  and  641).  Sutural  line  is  not  straight,  the  ridge  being 
fairly  distinct.  The  striations  on  the  shell  vary  to  a  considerable  extent 
(Figs.  634  to  637  and  639).  Four  polar  capsules  at  the  anterior  end, 
mostly  unequal  in  size  and  shape.  The  coiled  polar  filament  is  invisible 
in  formol  material.  Potassium  hydrate  solution  does  not  cause  its  extru- 
sion in  the  preserved  spores.  The  sporoplasm  is  finely  granular  with  two 
nuclei.  Dimensions  of  unstained  preserved  spores:  diameter  7.5  to  9//, 
polar  capsule  3  by  2.5/*. 

Remarks :  The  writer  was  able  to  study  forty  specimens  of  gall-bladder 
of  Alaskan  fishes,  chiefly  of  salmon,  which  have  been  collected  by  Professor 
Henry  B.  Ward,  during  the  summer  of  1919,  for  which  he  wishes  to  express 
his  deepest  appreciation.  The  examination  of  these  specimens  showed 
that  myxosporidia  were  found  only  in  one  of  the  gall-bladders,  and  that 
specimen  presented  a  fairly  heavy  infection  of  the  present  species. 

Family     SPHAEROSPORIDAE     Davis 
1917        Sphaerosporidae  Davis  1917  :  219 

The  characters  of  the  family  are  described  on  page  57. 


100  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [338 

Genus     SPHAEROSPORA    Thelohan 
1892        Sphaerospora  Th61ohan  1892  :  167 

The  characters  of  the  genus  are  described  on  page  57. 
Type  species:  Sphaerospora  diver  gens  Thelohan. 

SPHAEROSPORA  DIVERGENS  Thelohan 
[Figs.  183  to  186] 
1895        Sphaerospora  divergens  Th61ohan  1895  :  339-340 

1912        Sphaerospora  divergens  Parisi  1912  :  289 

1912        sphaerospora  divergens  Auerbach  1912  :  41-42 

Habitat:  Urinary  tubules  of  kidney  of  Blennius  phoUs  L.,  Crenilabrus 
melops  L.,  C.  pavo  Cuv.  et  V,  and  urinary  bladder  of  Hippoglossoides 
Hmandoides;  Concarneau,  Roscoff,  Napoli  (July),  Smalfjorden. 

Vegetative  form:  Rounded  discoidal  or  spherical  or  more  or  less 
elongate.  Ectoplasm  transparent,  without  real  pseudopodium.  Move- 
ments extremely  slow.  Endoplasm,  granular,  contains  fat  globules  and 
small  yellowish  granules.  Size  of  sporulating  individuals:  65/i  by  55/*, 
60)u  by  25/x,  60ju  by  20/i,  etc.  Polysporous  (Thelohan);  monosporous,  and 
disporous  (Auerbach). 

Spore:  Spherical.  Shell  with  fine  striations.  Two  polar  capsules 
divergent;  coiled  polar  filament  visible  in  fresh  state.  Sporoplasm  fills  the 
extracapsular  cavity  of  the  spore.  Dimensions:  lO/x  in  diameter,  often 
10/i  by  12/x,  the  larger  diameter  coinciding  with  sutural  plane,  thickness 
Bn  (Auerbach),  polar  capsules  about  4/li  long,  length  of  polar  filament 
20  to  25m. 

SPHAEROSPORA  ELEGANS  Thelohan 
[Figs.  187  and  188] 


1890 

Thdohan 

1890 

193-209 

1892 

Sphaerospora  elegans 

Thelohan 

1892 

167-175 

1894 

Chloromyxum  (Sphaerospora) 

elegans 

Gurley 

1894 

266 

1895 

Sphaerospora  elegans 

Th61ohan 

1895 

338-339 

1909 

Sphaerospora  elegans* 

Auerbach 

1909a 

:71 

1912 

Sphaerospora  elegans 

Parisi 

1912  : 

289 

Habitat:  Renal  tubules  of  kidney,  connective  tissue  of  ovary  and 
urinary  bladder  of  Gasterosieus  aculeatus  L.,  G.  pungitus  L.,  Loia  vulgaris 
Cuv.,    Phoxinus  laevis   L.;   Paris,   Bretagne,    Karlsruhe,    Lake   Garda. 

Vegetative  form:  Rounded  or  sUghtly  elongated,  not  exceeding  20  to 
25/Lt  in  diameter.  Protoplasm  homogeneous,  very  finely  granular,  contains 
numerous  refractive  globules,  probably  of  fatty  nature.  Pseudopodia 
lobose.    Movements  slow.     Disporous. 

*Misprinted  as  Sphaeromyxa  elegans. 


339]  STUDIES  ON  MYXOSPORIDIA—KUDO  101 

Spore:  Spherical,  somewhat  attenuated  at  the  anterior  end.  Sutural 
ridge  present,  terminating  in  a  small  projection  at  each  end  of  the  spore. 
Two  polar  capsules  spherical.  Coiled  polar  filament  not  visible  in  fresh 
state.    Dimensions:  diameter  lO^t  in  average,  sutural  diameter  about  ll^i. 

SPHAEROSPORA  ROSTRATA  Thelohan 
[Fig.  189] 
1895        Sphaerospora  rostrata  TWlohan  1895  :  339 

Habitat:  Malpighian   bodies   of  kidney   of   Mugil   sp.;   Roscoff,   Le 
Croisic,  Le  Vivier-sur-mer,  Marseille,  Banyuls. 
Vegetative  form:  Not  described. 

Sphore:  Subspherical.  Shell  shows  deep  longitudinal  striations  which 
end  in  sharp  spinous  edges  at  the  posterior  end.  Sutural  ridge  well 
marked.  Anterior  part  shows  enlargement  of  quadrangular  lamella,  which 
is  spinous  in  side  view.  Dimensions:  10  to  12/i  in  diameter,  sutural 
diameter  1  to  2/i  longer,  length  of  polar  filament  40/z. 

Remarks:  The  parasites  cause  the  degeneration  of  the  Malpighian 
bodies. 

SPHAEROSPORA  MASOVICA  Cohn 

[Figs.  190  to  192] 

1902        Sphaerospora  tnasovka  Cohn  1902  :  628-632 

Habitat:  Gall-bladder  of  Abramis  brama  L.;  Mauersee. 

Vegetative  form:  Polymorphous,  due  to  active  movements.  Trans- 
parent and  colorless,  while  in  motion.  Endoplasm  highly  granular,  contains 
yellowish  enclosures.  Ectoplasm  hyaline,  forms  a  narrow  layer  around  the 
body,  occasionally  developing  into  a  blunt  lobose  pseudopodium.  Pseudo- 
podia  of  two  kinds;  lobose  and  filose,  also  intermediate  forms.  Filiform 
pseudopodia  are  formed  and  retracted  more  slowly  than  the  lobose. 
Plasmotomy  is  of  probable  occurrence.  Two  spores  are  formed  in  each 
pansporoblast.  Size  variable:  lOfj.  (with  no  spore),  18/i  (with  sporob lasts), 
29^  (with  4  sporoblasts),  38/i  (with  22  sporoblasts).  Disporous(?),  poly- 
sporous. 

Spore:  Spherical.  Sutural  ridge  well  marked.  Polar  capsules  and 
sporoplasm  are  comparatively  small,  the  former  convergent.  By  warming 
the  spore,  polar  filament  is  extruded  and  at  the  same  time  two  filaments 
("starren  Faden")  are  made  visible  at  the  anterior  part  of  the  sutural  plane. 
Sporoplasm  with  two  nuclei,  no  vacuole  being  present.  Dimensions: 
diameter  8/i,  length  of  polar  filament  38/f,  length  of  sutural  filament  14/i. 

Remarks:  Cohn  did  not  observe  free  spores  in  the  gall-bladder.  He, 
however,  saw  many  free  spores,  separated  from  each  other,  in  the  intestine, 
concluding  that  the  body  and  pansporoblast  membrane  of  trophozoites, 
are  destroyed  in  the  intestine,  setting  the  spores  free. 


102  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [340 

SPHAEROSPORA  PLATESSAE  Woodcock 

[Figs.  193  and  194] 
1904        Sphaerospora  platessae  Woodcock  1904  :  59H50 

Habitat:  Otic-capsule  of  Pleuronectes  platessa  L.;  England. 

Vegetative  form:  Cysts  opaque  masses  about  1mm.  in  diameter. 
The  cartilage  was  greatly  hypertrophied.    Polysporous  (presumably). 

Spore:  Spherical.  Shell  unornamented.  Two  polar  capsules.  Sporo- 
plasm  with  several  refractive  granules,  but  without  any  vacuole.  Dimen- 
sions: diameter  8  to  9fi,  length  of  polar  filament  about  70/i. 

Remarks:  Woodcock  placed  this  species  provisionally  in  the  genus 
as  he  could  not  examine  any  fresh  material,  but  had  studied  smears  only. 

SPHAEROSPORA  ANGULATA  Fujita 

[Figs.  195  to  197] 

1912        Sphaerospora  angulata  Fujita  1912  :  261-262 

Habitat:  Kidney  of  Cyprinus  carpio  L.,  Carassius  auratus  L.;  Sapporo 
(Nippon). 

Vegetative  form:  Only  description:  "The  number  of  the  spore  in  the 
sporoblast  is  in  this  case  always  less  than  in  the  others,  rarely  exceeding 
two." 

Spore:  Somewhat  triangular,  with  convex  sides,  oval  in  sideview. 
Slightly  pointed  at  the  mid-posterior  margin  of  the  spore.  Shell  very  thin, 
faintly  marked  with  concentric  striations.  Two  oblong  polar  capsules 
are  of  unequal  size.  Dimensions:  length  7  to  8ju,  breadth  6  to  7^,  thickness 
5/x,  length  of  largest  polar  capsule  3.8/i,  length  of  polar  filament  twice  as 
long  as  that  of  the  spore. 

SPHAEROSPORA  POLYMORPHA  Davis 

[Figs.  198  and  199] 

1917        Sphaerospora  polymorpha  Davis  1917  :  231-232 

Habitat:  Urinary  bladder  of  Opsanus  tau;  Beaufort  (June,  July). 

Vegetative  form:  Elongate,  but  never  very  irregular  in  shape.  Slowly 
ameboid.  Body  colorless.  Ectoplasm  clearly  seen  in  younger  forms, 
forming  one  to  several  large  lobate  pseudopodia,  which  in  turn  extrude 
several  short,  conical  pseudopodia.  In  larger  forms,  ectoplasm  is,  often, 
recognizable  only  at  ends  of  pseudopodia,  which  in  such  cases  are  composed 
chiefly  of  endoplasm.  Endoplasm  granular,  vacuolated  in  some  smaller 
forms,  but  in  larger  individuals  vacuoles  are  indistinct  or  absent;  small  fat 
globules  abundant  in  large  forms;  numbers  of  rounded  sporoblast  cells 
can  be  distinctly  seen.  Size  of  large  trophozoites  35n  by  50/x.  Disporous 
and  polysporous  (polysporous  forms  rarely  contain  many  spores  at  the 
same  time). 


341]  STUDIES  ON  MYXOSPORJDIA— KUDO  103 

Spore:  Spherical,  sometimes  slightly  compressed  infero-superiorly. 
Sutural  ridge ;  on  each  side  are  a  number  of  concentric  striations  extending 
around  each  valve  parallel  to  sutural  line.  Polar  capsules  pyriform  and 
large.  Coiled  polar  filaments  indistinct.  Sporoplasm  finely  granular. 
Dimensions:  diameter  7  to  10/x,  (8/i  in  average),  polar  capsules  4  to  S/t  by 
2  to  2.5^. 

SPHAEROSPORA  sp.  Davis 

1917  Sphaerospora  sp.  Davis  1917  :  213 

Habitat:  Urinary  bladder  of  Lepisosteus  platystomus;  Gainesville,  Fla. 
Vegetative  form:  No  description. 
Spore:  Not  described. 

SPHAEROSPORA  sp.  Southwell  et  Prashad 

1918  Sphaerospora  sp.  Southwell  and 

Prashad  1918  :  347-348    . 

Habitat:  Under  the  scales  of  Barilius  barna;  from  the  vicinity  of  the 
Ruby  Mines,  Burma  (June). 

Vegetative  form :  The  cysts  occurred  in  very  large  numbers,  one  under 
each  scale. 

Spore:  Authors'  description:  "The  poor  condition  of  the  material  did 
not  allow  of  a  complete  account  of  its  structure,  but  the  bicapsulate, 
rounded  structure  of  its  spores  places  it  undoubtedly  in  the  genus  Sphaero- 
spora Thelohan." 

SPHAEROSPORA  CARASSII  nov.  spec. 
[Figs.  200  to  204] 

Habitat:  Gill  filament  of  Carassius  carassius  L.;  Tokio  (February). 

Vegetative  form:  Trophozoites  small  ameboid  in  groups  or  in  diffused 
condition  in  the  connective  tissue  of  the  gill  filament.  No  cyst  formation. 
The  number  of  trophozoites  in  groups  is  generally  small.  The  largest 
group  found  in  sections  was  96/i  by  36/n,  the  macroscopical  examination 
always  failing  to  trace  the  parasites.  The  trophozoites,  10  to  20/i  long, 
with  poorly  differentiated  protoplasm  and  usually  reticular  endoplasm 
without  any  particular  enclosure  (Fig.  200).  Ameboid  movements  not 
observed.  Schizogonic  multiplication  rapid,  each  of  the  daughter  indivi- 
duals developing  into  two  spores.  Disporous.  Other  sporous  characters 
could  not  be  determined. 

Spore:  Spherical  in  front  and  side  views,  tho  form  variable  to  some 
extent  (Figs.  201-203).  Shell  smooth.  Sutural  ridge  fairly  distinct.  Two 
polar  capsules,  broadly  pyriform,  of  equal  size  and  convergent,  located  at 


104  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [342 

the  anterior  end,  one  on  each  side  of  the  sutural  plane.  Coiled  polar 
filament  highly  distinct  (5  to  6  times)  in  vivo.  Sporoplasm  granular,  shows 
two  nuclei  when  stained;  no  vacuole  of  any  nature.  Dimensions  in  vivo: 
diameter  8  to  13/i,  polar  capsules  4  to  5^  by  2.5  to  3.5/x,  length  of  polar 
filament  35  to  40/x  (KOH  or  pressure). 

Remarks:  No  species  of  the  genus,  has  ever  been  found  in  the  branchiae. 
The  characters  of  the  spore,  however,  compel  the  writer  to  place  the 
form  in  the  present  genus. 

Genus    SINUOLINEA    Davis 
1917        Sinuolinea  Davis  1917  :  219 

The  characters  of  the  genus  are  described  on  page  57. 
Type  species:  Sinuolinea  dimorpha  Davis. 

SINUOLINEA  DIMORPHA  Davis 
[Figs.  205  to  213] 

1916  Sphaerospora  dimorpha  Davis  1916  :  333-377 

1917  Sinuolinea  dimorpha  Davis  1917  :  232-233 

Habitat:  Urinary  bladder  and  ureter  of  Cynoscion  regaJis;  Beaufort. 

Vegetative  form:  Disporous  and  polysporous  trophozoites  differ 
distinctly  from  each  other.  Disporous  trophozoites  irregular,  colorless, 
transparent  and  show  slow  movements.  When  attached  to  the  epithelium, 
rounded  with  one  to  several  pseudopodia.  Differentiation  of  protoplasm 
distinct.  Occasionally  endoplasm  contains  one  or  more  erythrocytes. 
Average  diameter  of  full-grown  form  25  to  30//. 

Polysporous  form:  when  attached  to  the  bladder  epithelium,  the  free  end 
is  drawn  out  into  a  long,  cylindrical  process,  covered  with  numerous  short, 
hairlike  ectoplasmic  processes.  While  not  movable,  these  processes  are 
readily  absorbed  and  reformed.  When  the  trophozoite  is  detached  from 
the  epithelium,  the  larger  end  gives  rise  to  numerous  conical  or  arborescent 
pseudopodia,  by  means  of  which  the  trophozoite  moves  slowly.  Endoplasm 
extends  into  the  proximal  portion  of  large  pseudopodia.  It  is  granular 
and  vacuolated,  contains  numerous  fat  globules,  refractive  granules, 
yellowish  crystals  (hematoidin?)  and  erythrocytes  in  various  stages  of 
disintegration.  Endoplasm  also  contains  gemmules,  each  composed  of 
outer  layer  and  finely  granular  central  portion.  Size  varies  greatly:  up  to 
575m  by  90m. 

Spore:  Spherical.  Sutural  ridge  well  marked.  Polar  capsules  large 
and  spherical.  Sporoplasm  forms  a  rounded  granular  mass.  Dimensions: 
diameter  15m,  diameter  of  polar  capsules  4.5m,  length  of  polar  filament 
27  to  35m. 


3431  STUDIES  ON  MYXOSPORJDJA—KUDO  105 

SINUOLINEA  CAPSULARIS  Davis 
[Figs.  214  to  216] 
1917        Sinuolinea  capstdaris  Davis  1917  :  233 

Habitat:  Urinary  bladder  of  Paralichthys  albiguttus,  P.  dentatus, 
Spheroides  maculatus;  Beaufort  (July,  August). 

Vegetative  form :  Rounded  to  irregular  shape.  Body  colorless  or  light 
yellow.  Progressive  movements  slow.  Pseudopodia  large  branched  or 
arborescent,  formed  entirely  of  ectoplasm.  Ectoplasm  transparent  and 
usually  granular,  merging  gradually  with  the  endoplasm.  Endoplasm 
contains  numberous  fat  globules.  In  large  trophozoites,  gemmules  are 
observed.  The  gemmules  are  more  finely  granular  and  more  transparent 
than  the  surrounding  protoplasm  and  are  practically  identical  with  the 
small,  free  trophozoites.  Trophozoites  containing  several  gemmules  are 
usually  rounded  and  motionless  and  appear  to  be  more  or  less  degenerate. 
Disintegration  of  such  trophozoites  were  actually  observed.  Sporulating 
trophozoites  were  rare  and  were  never  seen  to  contain  gemmules.  Size  up 
to  40m  in  diameter.    Disporous  and  polysporous(?). 

Spore:  Spherical,  sometimes  slightly  elongated.  Sutural  plane  much 
twisted  on  its  axis.  Sutural  ridge  very  distinct.  Polar  capsules  and  cap- 
sulogenous  cells  large  occupying  more  than  one-half  of  the  cavity  of  spore. 
Coiled  polar  filament  distinct,  Sporoplasm  granular  contains  numerous 
fat  globules.  Dimensions:  diameter  12  to  14/i,  diameter  of  polar  capsules 
4.5iu,  length  of  polar  filament  SOyit. 

SINUOLINEA  ARBORESCENS  Davis 
[Figs.  217  and  218] 
1917        Sinuolinea  arbor escens  Davis  1917  :  233 

Habitat:  Urinary  bladder  of  Siphostonia  floridae;  Beaufort 
Vegetative  form:  Rounded  or  irregular.  Body  colorless  or  light  yellow. 
Actively  ameboid,  forming  large  arborescent  pseudopodia  of  ectoplasm. 
Ectoplasm  well  developed,  hyaline  and  homogeneous.  Endoplasm  coarsely 
granular,  sometimes  containing  a  few  fat  globules.  Larger  trophozoites 
are  less  active  and  the  ectoplasm  less  distinct.  In  sporulating  trophozoites 
the  ectoplasm  may  entirely  disappear,  the  entire  trophozoite  consisting  of 
a  coarsely  granular  mass.  Diameter  of  rounded  sporulating  trophozoites 
75ju.     Polysporous. 

Spore:  Rounded,  in  front  view,  slightly  elongated  in  the  anterior  end 
view.  Polar  capsules  large.  Sutural  ridge  prominent,  makes  a  character- 
istic S-shaped  turn  on  the  anterior  end.  Coiled  polar  filaments  distinct. 
Dimensions:  length  15fx,  breadth  12/i,  diameter  of  polar  capsules  5/i. 


106  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [344 

SINUOLINEA  OPACITA  Davis 
[Fig.  219] 
1917        Sinuolinea  opacita  Davis  1917  :  234 

Habitat:  Urinary  bladder  of  Paralichihys  albiguttus;  Beaufort  (August). 

Vegetative  form:  Rounded  or  slightly  irregular.  Body  colorless  and 
opaque.  Movements  slow.  Pseudopodia  short  lobose.  Ectoplasm  not 
distinct,  except  around  ends  of  pseudopodia,  where  it  forms  a  thin  hyaline 
layer.  Endoplasm  opaque,  finely  granular,  with  numerous  greenish-yellow 
fat  globules  varying  greatly  in  size.  Diameter  of  rounded  sporulating 
trophozoites  22/Lt,  exceptionally  large  trophozoites  100/x.    Disporous. 

Spore:  Nearly  spherical,  with  flattened,  lateral  appendages  extending 
from  the  posterior  side.  Sutural  plane  slightly  twisted  on  its  axis.  Sutural 
ridge  distinct.  Polar  capsules  large.  Coiled  polar  filament  distinct. 
Sporoplasm  finely  granular,  containing  several  comparatively  large  fat 
globules.    Dimensions:  diameter  12  to  13/x,  diameter  of  polar  capsules  ^n. 

SINUOLINEA  BRACHIOPHORA  Davis 
[Fig.  220] 
1917        Sinuolinea  brachiophora  Davis  1917  :  234 

Habitat:  Urinary  bladder  of  Paralichthys  albiguttus;  Beaufort  (August 
only  in  one  fish). 

Vegetative  form:  Rounded  to  somewhat  irregular.  Body  colorless. 
Ectoplasm  hyaline.  Endoplasm  granular,  with  numerous  large  fat  glob- 
ules.    Disporous. 

Spore:  Nearly  spherical,  with  a  long  lateral  appendage  from  each 
valve.  These  appendages  are  empty  except  at  extreme  distal  end,  which 
contains  a  granular  mass,  probably  the  remains  of  the  parietal  cell.  Sutural 
plane  slightly  oblique  to  longitudinal  axis.  Sutural  ridge  distinct.  Polar 
capsules  and  capsulogenous  cells  large,  occupying  more  than  half  of  cavity 
of  spore.  Sporoplasm  finely  granular.  Dimensions:  length  exclusive  of 
appendages  9  to  ll/x,  length  of  appendages  18  to  22/i,  breadth  of  spore  9/i, 
diameter  of  polar  capsules  3.5/*. 

Remarks:  Davis  mentions  that  in  many  respects  this  species  is  very 
similar  to  S.  opacita,  which  occurs  in  the  same  host. 

Suborder     PLATYSPOREA     nom.  nov. 

The  definition  of  the  suborder  is  recorded  on  page  57. 

Family     MYXIDIIDAE     Thelohan 

1892  Myxididees  Thaohan  1892  :  173,  175 

1893  Myxidiidae  Gurley  1893  :  412 

The  characters  of  the  family  are  described  on  page  57. 


345] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


107 


Genus     MYXIDIUM     Biitschli 
1882        Myxidium  Butschli  1882  :  PI.  38 

The  characters  of  the  genus  are  described  on  page  58. 
Type  species:  Myxidium  lieberkiihni  Butschli. 


MYXIDIUM  LIEBERKtJHNI  Butschli 
[Figs.  221  to  240] 


1854 

1879 

1881 

1882 

Myxidium 

1883 

1891 

Myxidium 

1894 

Myxidium 

1895 

Myxidium 

1895 

Myxidium 

1898 

Myxidium 

1902 

Myxidium 

1902 

Myxidium 

1906 

Myxidium 

1909 

Myxidium 

1912 

Myxidium 

1912 

Myxidium 

1916 

Myxidium 

1916 

Myxidium 

lieberkuhni 
lieberkuhni 
lieberkuhni 
lieberkuhni 
lieberkiihni 
lieberkuhni 
lieberkuhni 

lieberkuhni 
lieberkuhni 
lieberkuhni 
lieberkuhni 
lieberkuhni 
lieberkuhni 


Lieberkuhn 

1854 

5H5,349 

Leuckart 

1879 

246 

Butschli 

1881 

638-648 

Butschli 

1882 

593-595 

Balbiani 

1883 

201-202,  274-275 

Pfeiffer 

1891 

20,  91,  105,  127 

Gurley 

1894 

283-289 

Thelohan 

1895 

340 

Cohn 

1895 

5-36 

Doflein 

1898 

229,  341 

Prenant 

1902a 

:  200-217 

Laveran  and 

Mesnil 

1902 

469-472 

L^ger  and  Hesse  1906 

720 

Auerbach 

1909a 

:71 

Schroder 

1912 

326-327 

Parisi 

1912 

286 

Mavor 

1916a 

:66-68 

Mavor 

1916b 

:  373-378 

Habitat:  Urinary  bladder  of  Esox  lucius  L.,  Lota  lota  L.  (L.  vulgaris) \ 
France,  Canada  (Georgian  Bay),  U.  S.  A.,  (Wisconsin,  Lake  Mendota), 
Italy  (Lago  Maggiore,  Lago  di  Como,  Milano),  Germany. 

Vegetative  form:  Form  variable  with  lobose  or  immovable  filiform 
pseudopodia.  Clear  differentiation  of  protoplasm.  Cohn  described  third 
layer  of  protoplasm  (mesoplasm).  Endoplasm  yellowish  in  older  tropho- 
zoites, contains  yellow  globules,  fat  globules  and  hematoidin  crystals. 
Size  varying  with  age  up  to  a  maximum  length  of  300/i  by  a  breadth  of 
136/x  (Butschli).  Plasmotomous  multiplication  active.  Cohn  described 
budding  of  larger  forms,  while  Laveran  et  Mesnil  observed  only  the  division 
of  smaller  forms.  Each  pansporoblast  develops  into  two  spores.  Poly- 
sporous. 

Spore:  Elongated  fusiform.  Shell  with  longitudinal  striations.  Polar 
capsule  at  each  end  of  the  spore.  The  longer  axis  of  polar  capsules  coincides 
with  that  of  spore.  Dimensions:  length  18  to  20/i,  width  5  to  6At.  Mavor's 
measurement:  polar  capsules  5/i  by  2.5  to  Zn,  length  of  polar  filament 
40  to  45/i. 


106  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [346 

MYXIDIUM  INCURVATUM  Thelohan 
[Figs.  241  to  251] 


1892 

Myxidium  ?  inciirvatum 

Th61ohan 

1892a 

:  1093-1094 

1895 

Myxidium  incurvatum 

ThflohaTi 

1895 

341 

1912 

Myxidium  incurvatum 

Parisi 

1912 

286-287 

1912 

Myxidium  incurvatum 

Auerbach 

1912 

4,39 

1916 

Myxidium  incurvatum 

Georg6vitch 

1916 

90-91 

1917 

Myxidium  incurvatum 

Davis 

1917 

234^235 

Habitat:  Gall-bladder  of  Xerophis  aequoreus  L.,  N.  annulatus,  N. 
lumbriciformis,  Blennius  pholis  L.,  Callionymus  lyra  L.,  Fundulus  majalis, 
Gambusia  affinis,  Hippocampus  brevirostris,  Mugil  cephalus,  Scorpaena 
scrofa  L.,  Syngnathus  acus  L.,  S.  typhle;  Roscofif,  Concarneau,  Marseille, 
Banyuls,  Napoli,  Bergen,  Monaco,  Beaufort  (July). 

Vegetative  form:  Thelohan  describes  as  follows:  Trophozoites  usually 
small,  sometimes  reaching  a  considerable  size.  Pseudopodia  lobose. 
Protoplasm  pale  and  finely  granular  with  refractive  globules.  Disporous. 
According  to  Parisi  and  Davis  rarely  monosporous.  Georgevitch  observed 
apparently  the  polysporous  form. 

Parisi's  form:  ectoplasm  hyaline,  endoplasm  granular.  Monosporous 
form  25/1  long. 

Davis's  form:  lobose  pseudopodia,  occasionally  being  drawn  out  into  a 
long  process.  Many  trophozoites  often  cling  together  closely.  Diameter 
of  rounded  disporous  forms  about  13  to  15m,  that  of  monosporous  forms 
about  10  to  ll/t. 

Spore:  Thelohan's  description  is  as  follows:  Irregular  fusiform.  Long- 
est axis  curved  into  S-form,  both  ends  sharply  pointed  and  directed  toward 
opposite  directions.  Polar  capsule  opening  on  opposite  side  of  the  spore, 
in  some  spores  the  axis  of  the  polar  capsules  being  parallel  to  each  other. 
Dimensions:  length  8  to  9/x,  breadth  4  to  5/iz,  length  of  polar  filament  10 
to  15m. 

Parisi  gave  the  following  dimensions:  length  10  to  12/x,  breadth  5  to 
6/t,  length  of  polar  capsule  3m,  length  of  polar  filament  28m. 

According  to  Georgevitch,  young  spores  are  not  curved  (Fig.  245). 

Davis's  form;  Polar  filaments  when  extruded  in  HCl  remained  tightly 
coiled.  Dimensions:  length  8  to  9m,  Avidth  5  to  6m,  diameter  of  polar 
capsule  about  3>n. 

Remarks:  As  are  shown  in  figures,  Davis's  form  seems  to  be  some- 
what different  from  the  European  forms. 


347]  STUDIES  ON  MYXOSPORIDIA—KUDO  109 

MYXIDIUM  SPHAERICUM  Thelohan 
[Fig.  252] 
1895        Myxidiunt  sphericum  (corr. 

sphaericum)  Thelohan  1895  :  341-342 

Habitat:  Gall-bladder  of  Belone  acus  (Belone  belone  L.);  Banyuls, 
Le  Vivier-sur-Mer. 

Vegetative  form :  Trophozoites  spherical  or  subspherical,  not  exceeding 
20  to  22)Li  in  diameter  with  lobose  pseudopodia  formed  from  the  entire 
surface.  Endoplasm  granular,  contains  small  refractive  granules.  Dis- 
porous. 

Spore:  Form  similar  to  M.  incurvatum,  but  much  greater.  Coiled  polar 
filament  distinctly  visible  in  fresh  spore.  Dimensions:  length  15  to  20/t, 
width  7  to  8m,  length  of  polar  filament  60/i  (KOH). 

MYXIDIUM  HISTOPHILUM  Thelohan 
[Fig.  253] 
1895        Myxidiunt  histophilum  Th61ohan  1895  :  341 

Habitat:  Connective  tissue  of  kidney  and  ovary  of  Leuciscus  phoxinus 
L.  (Phoxinus  laevis  Ag.);  France. 

Vegetative  form:  Small  mass. 

Spore:  Fusiform,  being  compressed  at  the  middle  part.  Shell  with 
longitudinal  striations.    Length  of  the  spore  15/i. 

MYXIDIUM  sp.  Gurley 


[Fig.  254] 

1851 

Leydig 

1851 

:  226,  234 

1852 

Leuckart 

1852 

:436 

1894 

Myxidiunt  ?  sp.  incert.'             Gurley 

1894  : 

:290 

1899 

Myxidiunt  sp.                            Labb^ 

1899  : 

:92 

Habitat:  Gall-duct  of  Raja  hatis  L. 
Vegetative  form:  No  description. 
Spore:  Not  described.    One  figure. 

MYXIDIUM  DANILEWSKYI  Laveran 
[Figs.  255  to  257] 

1887  DanUewsky         1887  :  35 

1897  Myxidiunt  danilewskyi  Laveran  1897  :  725-726 

1898  Myxidiunt  danilewskyi  Laveran  1898  :  27-30 

Habitat:  Kidney  of  Emys  orbicularis  L.;  France. 

Vegetative  form:  Form  elongated,  circular  in  cross-section,  tapering 
toward  the  ends.  Body  of  greenish  color,  occupying  the  lumen  of  the  renal 
tubules  of  the  kidney.    Body  bent  along  the  cavity  of  the  tubule.    Endo- 


no  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [348 

plasm  granular,  ectoplasm  covering  the  entire  surface  of  the  body  as  a 
thin  layer.    Each  pansporoblast  develops  two  spores.    Polysporous. 

Spore :  Elongated  fusiform,  similar  to  M.  lieherkuhni,  but  much  smaller. 
Polar  capsule  at  each  end,  extrudes  filament  under  the  action  of  nitric 
acid.  Sporoplasm  granular  with  one  nucleus.  Dimensions:  length  12/*, 
breadth  3  to  4/i. 

MYXIDIUM  GIGANTEUM  Doflein 

[Fig.  258] 

1898        Myxidium  giganteum  Doflem  1898  :  285-286 

Habitat:  Gall-bladder  of  Raja  asterias;  Napoli. 

Vegetative  form:  Rounded  trophozoites.  Lobose  pseudopodia  with 
slow  movement,  show  remarkable  dimensions.  Posterior  portion  forms 
"Stemm-pseudopodien."  Small  form  club-shaped.  Endoplasm  is  of 
yellowish  color.  Diameter  of  large  form  500/x,  of  medium  sized  200/i,  small 
individuals  70-90^t,  quite  young  ones,  polymorphous  8  to  40/x.  Larger 
individual  up  to  700/i  by  180/i.  Many  trophozoites  form  a  cyst-like  motion- 
less stage,  in  which  many  individuals  seem  to  be  covered  with  a  common 
gelatinous  envelope.    Each  pansporoblast  forms  two  spores.    Polysporous. 

Spore:  Elongated.  Fusiform  in  front  view;  in  side  view,  one  valve 
arch-form,  the  other  being  flat.  Transparent.  Two  polar  capsules,  one 
at  each  end.  Coiled  polar  filament  is  clearly  seen  in  larger  polar  capsules. 
Dimensions :  length  28ju,  breadth  8/t,  polar  capsules  8/x  by  4/1. 

MYXIDIUM  BARBATULAE  Cepede 

1906        Myxidium  barbatulae  C6pSde  1906  :  67 

1906        Myxidium  barbatulae  Cepede  1906a  :  15-16 

Habitat:  Kidney  of  Cobitis  barbatula  L.;  Isere. 

Vegetative  form:  Trophozoites  form  cysts.  Form  and  size  vary 
greatly.    Average  size:  400  to  500m  in  length  and  200/x  in  breadth. 

Spore:  Irregular  fusiform.  Polar  capsule  at  each  end  of  the  spore. 
Shell  longitudinally  striated,  number  being  variable.  Dimensions:  length 
12  to  15/i,  breadth  about  6/x,  polar  capsules  5/i  by  2.5  to  Six. 

MYXIDIUM  GIARDI  Cepede 
[Figs.  259  to  261] 
1906       Myxidium  giardi  C6pede  1906a  :  16;  1906b  :  170-173 

1908        Myxidium  giardi  C6pSde  1908    :  93-95 

1908        Myxidium  giardi  C6pede  1908a  :  8 

Habitat:  Kidney  of  Anguilla  vulgaris  Flem.;  Wimereux  (August). 

Vegetative  form:  Subspherical  white  cysts,  800  to  900/x  in  diameter, 
surrounded  by  a  thick  (up  to  30/x)  membrane,  composed  of  the  connective 
tissue  of  the  host. 


349]  STUDIES  ON  MYXOSPORIDIA—KUDO  111 

Spore:  Irregular  fusiform,  greatly  enlarged  at  the  middle  portion. 
Plane  of  symmetry  of  the  spore  coincides  with  the  sutural  plane.  Shell 
thick  with  9  to  11  longitudinal  striations  on  each  valve,  which  are  more 
clearly  seen  on  spores  stained  with  iron  hematoxylin.  Polar  capsule  at 
each  end.  Coiled  polar  filament  distinct.  Sporoplasm  finely  granular 
with  two  nuclei  and  refringent  globules.  Dimensions  in  vivo:  length  9  to 
lO^t,  width  5  to  5.6/1,  thickness  4.75  to  5/x,  polar  capsules  3. 5m  by  2ju. 

MYXIDIUM  PFEIFFERI  Auerbach 

[Figs.  262  to  265] 

1908        Myxidium  pfeiferi  Auerbach  1908  :  459-464 

1910        Myxidium  pfeiferi  Auerbach  1910c  :  171-172 

Habitat:  Gall-bladder  of  Tinea  vulgaris  Cuv.;  Karlsruhe. 

Vegetative  form:  Observations  in  sections.  More  or  less  flattened, 
disc-form,  often  enrolled.  The  ectoplasm  finely  granular,  without  large 
pseudopodia.  It  is  not  usually  distinguishable  from  the  endoplasm  which 
is  highly  alveolar  and  contains  numerous  nuclei,  but  no  enclosures. 

Spore:  Form  varies  to  some  extent.  Similar  to  Myxidium  lieberkiihni; 
slightly  curved.  Shell  with  fine  longitudinal  striations.  Polar  capsules 
two,  one  at  each  end.  Polar  filament  is  extruded  by  adding  one  drop  of 
water  to  the  smear  of  the  spore,  which  had  been  dessicated  for  24  hours. 
Sporoplasm  with  one  or  two  nuclei,  in  one  case  with  four  small  nuclei, 
which  is  thought  to  be  an  abnormal.  Dimensions:  length  13  to  IS^i, 
breadth  5.2  to  5.8/x,  length  of  polar  capsule  5.2  to  6/i,  length  of  polar  fila- 
ment 45  to  54/i. 

MYXIDIUM  INFLATUM  Auerbach 
[Fig.  266] 


1909 

Myxidium  inflatum 

Auerbach 

1909  :  72-74 

1909 

Myxidium  inflatum 

Auerbach 

1909a  :  31 

1910 

Myxidium  inflatum 

Auerbach 

1910c  :  172 

1912 

Myxidium  inflatum 

Auerbach 

1912  :  39 

Habitat:  Gall-bladder  of  Cydopterus  lumpus  L.;  Bergen  (September). 

Vegetative  form:  Extremely  polymorphous.  Rounded,  spherical, 
or  much  elongated.  Ameboid  movements  very  active.  Differentiation  of 
protoplasm  is  sharp  and  clear,  which  is  best  observed  in  individuals  in 
motion;  highly  hyaline  ectoplasm  forms  very  long  lobose  pseudopodia, 
into  which  granular  endoplasm  flows  in  slowly.  Size  variable.  Rounded 
large  form  44  to  45/i  in  diameter.  Fully  grown  spores  are  set  free  from  the 
mother  trophozoite  in  comparatively  short  time.  Spore  formation  similar 
to  that  of  Myxidium  bergense.  Disporous  and  polysporous  (5  spores  in 
maximum). 


Keysselitz 

1908  : 

289 

?  Myxidium  sphaericum 

Auerbach 

1909  : 

75-76 

Myxidium  bergense 

Auerbach 

1910  : 

61 

Myxidium  bergense 

Auerbach 

1910c 

:172 

Myxidium  bergense 

Auerbach 

1912  : 

18-39 

Myxidium  bergense 

Mavor 

1915  : 

30-31 

jall-bladder  of  Gadus 

virens  L.,  G.  aet 

'lefinis 

,G.l 

112  JLUNOJS  BIOLOGICAL  MONOGRAPHS  [350 

Spore:  Very  broad  compared  with  the  length.  The  longitudinal  axis 
is  curved  in  S  shape.  Polar  capsule  situated  in  opposite  way  at  each  end 
of  the  spore.  Dimensions:  length  20.8  to  23.4pi,  breadth  13  to  15.6/i, 
polar  capsules  7.8^,  length  of  polar  filament  90  to  lOOjit  (KOH). 

MYXIDIUM  BERGENSE  Auerbach 
[Fig.  267] 

1908 
1909 
1910 
1910 
1912 
1915 

Habitat: 

Pleuronectes  platessa  and  Sehastes  viviparus,   Melanogrammus   aeglefinis; 
Norway  (Bergen),  Canada  (St.  Andrew,  July  to  September). 

Vegetative  form:  Rounded  or  elongated,  as  the  result  of  formation  of 
various  pseudopodia.  Trophozoites  partly  free,  partly  attached  to  the 
epithelium  of  the  bladder.  Size  up  to  54/x  in  diameter.  Pseudopodia  of 
two  kinds:  lobose  and  long  filose,  sometimes  slightly  branched.  Mavor 
observed  a  cyst-like  stage  under  certain  conditions,  which,  he  thinks,  may 
be  due  to  some  exceptional  conditions  of  the  parasite.  Plasmogamy. 
Monosporous,  disporous  and  polysporous. 

Spore:  Fusiform.  Main  axis  curved  into  S  shape.  Form,  roughly 
speaking,  very  much  similar  to  that  of  M.  sphaericum  Thel.  Dimensions: 
length  16.2  to  IQjli,  breadth  7  to  9n,  length  of  polar  capsules  5An,  length 
of  polar  filament  about  three  times  as  that  of  spore. 

MYXIDIUM  PROCERUM  Auerbach 
[Fig.  268] 
1910        Myxidium  procerum  Auerbach  1910  :  61-62 

1910        Myxidium  procerum  Auerbach  1910c  :  172-173 

1912        Myxidium  procerum  Auerbach  1912  : 4, 39 

Habitat:  Gall-bladder  of  Argentina  situs  As.;  Bergen. 

Vegetative  form:  Not  observed. 

Spore:  Greatly  elongated  and  narrow.  Sporoplasm  with  one  or  two 
nuclei.  Dimensions:  length  21.6  to  25.2/i,  breadth  3.6  to  4/x,  length  of 
polar  capsule  7.2/1. 

MYXIDIUM  MACKIEI  Bosanquet 

[Figs.  269  to  271] 

1910        Myxidium  mackiei  Bosanquet  1910  :  436-438 

Habitat:  Renal  tubules  of  kidney  of  Trionyx  ganzeticus;  Bombay. 
Observations  on  three  slides. 


351]  STUDIES  ON  MYXOSPORIDJA—KUDO  113 

Vegetative  form:  The  largest  trophozoite  160^  by  21  fi.  No  distinction 
between  ectoplasm  and  endoplasm  could  be  drawn,  except  in  a  few  indi- 
viduals in  which  there  was  a  cyst- wall.  Spores  are  formed  in  pairs.  Proto- 
plasm with  two  kinds  of  nuclei,  some  vesicular,  others  smaller  and  com- 
pact.   Polysporous. 

Spore:  Fusiform  with  rather  pointed  ends.  Shell  finely  striated.  Two 
comparatively  small  polar  capsules,  one  at  each  end.  Sporoplasm  with 
one  or  two  nuclei,  contains,  often,  two  large  vacuoles.  Dimensions: 
length  16/x  (a  few  11  ix),  breadth  5^  (many  broader  than  this). 

Remarks:  The  discoverer,  J.  P.  Mackie  mentioned  that  the  parasites 
did  not  appear  to  excite  any  reaction  in  the  tissue  of  the  host,  the  animal's 
health  being  unaffected. 

MYXIDIUM  MACROCAPSULARE  Auerbach 
[Figs.  272  and  273] 

1910        Myxidium  macrocapsulare         Auerbach  1910  :  440-441 

Habitat:  Gall-bladder  of  Scardinius  eryihrophihalmus  L.;  Karlsruhe. 

Vegetative  form:  Not  observed. 

Spore :  Elongated  elliptical  when  viewed  at  right  angles  to  sutural  plane. 
Shell  somewhat  thick  with  longitudinal  striations  parallel  to  the  sutural 
line.  In  side-view,  both  ends  pointed  in  diagonally  opposite  directions. 
Polar  capsules  are  comparatively  large,  one  at  each  end,  opening  at  the 
sharply  pointed  end.  Dimensions:  length  10  to  12/x,  breadth  6/x,  polar 
capsules  3  to  4^t. 

Remarks:  No  pathological  change.    Bile  was  clear. 


MYXIDIUM  sp.  Awe'rinzew 

[Figs.  274  to  276] 

1908 

Myxidium  sp.                            Awerinzew 

1908 

:  33,  43,  45,  55 

1909 

Myxidium  sp.                            Awerinzew 

1909 

:  76,  78,  80,  81 

1911 

Myxidium  sp.                            Awerinzew 

1911 

:  199-204 

Habitat:    Gall-bladder  of  Cottus  scorpius;    Aleksandrowsk,  North  Sea. 

Vegetative  form:  Trophozoites  are  small.  The  protoplasm  is  differ- 
entiated into  ectoplasm  and  endoplasm  in  some  specimens.  Very  active 
formation  of  filiform  pseudopodia  of  various  length.  Degenerating  troph- 
ozoites, with  one  or  two  empty  spaces  are  often  noticed.  Each  spore 
is  formed  independently  from  each  other.  Monosporous,  disporous  and 
polysporous  (with  three  spores). 

Spore:  Form  similar  to  Myxidium  incurvatum.  Young  spores  not 
curved.    Dimensions:  length  20  to  35^1,  breadth  10  to  15/i. 


114  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [352 

MYXIDIUM  DEPRESSUM  Parisi 

[Figs.  277  and  278] 
1912        Myxidium  depressum  Parisi  1912  :  287 

Habitat:  Gall-bladder  of  Citharus  Unguatula  Gthr.;    Napoli  (August). 

Vegetative  form:  Not  observed. 

Spore:  Fusiform  with  greatly  attenuated  extremities  in  front  view; 
flattened  and  curved  in  S-form  in  profile.  The  axis  of  polar  capsules 
parallel  to  each  other.  Coiled  polar  filament  visible  in  vivo.  Sporoplasm 
with  two  nuclei,  occupies  the  extracapsular  cavity  of  the  spore.  Dimen- 
sions: length  12  to  14/x,  breadth  5.5  to  6/x,  thickness  2.5  to  3^1,  polar  cap- 
sules 5.5  to  6/Li  by  2.3^1,  length  of  polar  filament  30m. 

MYXIDIUM  OVIFORME  Parisi 
[Figs.  279  and  280] 

1912        Myxidium  oviforme  Parisi  1912  :  287-288 

1912        Myxidium  oviforme  Auerbach  1912  :  39 

Habitat:  Gall-bladder  of  Apogon  rex  mullorum  Cuv.,  Coris  julis  Gthr., 
Gadus  callarias  L.,  Trutta  solar  L.;  Napoli  (August),  Norwegian  coast. 

Vegetative  form:  Unobserved  by  Parisi.  Auerbach's  observation  is  as 
follows : 

Trophozoites,  small  ameboid,  usually  spherical.  Size  10  to  12/i  in 
diameter.    Monosporous  (probably). 

Spore:  Oval  with  rounded  extremities,  slightly  pointed  at  the  foramina 
of  polar  capsules.  Shell  with  numerous  fine  striations  running  longitudin- 
ally. Polar  capsules  being  often  invisible,  opening  a  little  above  and  below 
of  the  hypothetical  horizontal  plane.  Sporoplasm  fills  the  extracapsular 
cavity  of  the  spore,  leaving  little  space  at  the  extremities  of  the  polar 
capsules.  Dimensions:  length  11/i,  breadth  8  to  8.5/i,  polar  capsules 
4.5)u  by  2>ix,  length  of  polar  filament  30  to  35/i.  Auerbach's  measurements: 
length  12  to  13/i,  breadth  8  to  9/i,  polar  capsules  about  4/t  long. 

MYXIDIUM  ANGUILLAE  Ishii 

[Figs.  281  to  284] 

1915        Myxidium  anguiUae  Ishii  1915  :  372-382 

Habitat:  Integument  of  the  side  of  the  body  of  Anguilla  japonic  a 
Temm.  et  Sch.;  Schizuoka,  Nippon  (October).  Number  of  the  cysts 
visible  to  unaided  eye,  10  and  9  on  the  left  and  the  right  side  respec- 
tively. 

Vegetative  form:  Trophozoites  form  white  and  sharply  contoured  cysts. 
Cysts,  spherical  or  oval,  surrounded  by  a  membranous  connective  tissue 
(about  2ix  thick)  of  the  host.  Protoplasm  is  clearly  differentiated  into 
ectoplasm  and  endoplasm.  Diffuse  infiltration  also  occurs.  Size  measured 
along  the  skin,  1.2  to  2mm.  in  diameter;  in  sections  1.174mm.  by  0.658mm. 


353]  STUDIES  ON  MYXOSPORIDIA—KUDO  115 

Spore:  Form  similar  to  Myxidium  pfeiferi  Auerbach,  but  rather 
straight  fusiform,  rarely  slightly  bent.  In  many  spores  the  shell  tapers 
to  a  sharp  point  at  each  end.  Shell  striated  longitudinally,  22  in  all 
(2  sutural  ridges?).  Two  polar  capsules,  one  at  each  end.  Sporoplasm 
usually  with  two  nuclei.  Dimensions:  length  9.1ai,  breadth  2,8^,  length 
of  polar  capsule  3.5/x. 

MYXIDIUM  sp.  Mavor 
[Figs.  285  and  286] 

1915  Myxidium  sp.  Mavor  1915  :  32 

Habitat:  Gall-bladder  of  Pseudopleuronectes  americanus ;  New  Bruns- 
wick (Canada),  of  rare  occurrence. 

Vegetative  form:  Observations  in  smears  are  as  follows:  Spheroidal, 
with  numerous  long  pseudopodia  on  one  side,  which  suggests  the  attach- 
ment of  the  trophozoite  to  the  bladder.  Trophozoites  without  any  spore. 
Pansporoblasts  spherical,  15  to  16/i  in  diameter. 

Spore:  Spindle  shaped.  The  long  axis  being  slighly  bent  in  S-form. 
Two  pear  shaped  polar  capsules,  one  at  each  end  of  spore.  Coiled  polar 
filament  visible  in  fresh  state.  Dimensions:  length  14  to  15/x,  breadth 
6  to  7.5/i,  polar  capsules  4/i  by  2.5/x,  length  of  polar  filament  90  to  95/i 
(ammonia  water). 

MYXIDIUM  GADI  Georgevitch 
[Figs.  287  to  290] 

1916  Myxidium  gadi  Georg6vitch         1916  :  88-89 

1917  Myxidium  gadi  Georg6vitch         1917c  :  797-799 
1919        Myxidium  gadi                        Georgevitch         1919  :  251-289 

Habitat:  Gall-bladder  of  Gadus  pollachius,  Solea  vulgaris  Quens; 
Roscoff  (September). 

Vegetative  form:  Highly  polymorphous.  Spherical  or  oval.  Large 
forms  fill  up  the  bladder.  Ectoplasm  hyaline  and  transparent,  forming 
one  long  or  many  short  lobose  pseudopodia.  Endoplasm  colorless  and 
finely  granular,  contains  more  or  less  large  numbers  of  nucleus.  Mono- 
sporous,  disporous  and  polysporous. 

Spore:  Fusiform  with  attenuated  ends.  Young  spores  more  attenuated 
than  the  fully  grown  forms.  The  main  axis  of  the  spore  coincides  with  the 
longitudinal  axis  of  the  polar  capsules,  with  slight  deviation.  Two  nuclei 
of  the  sporoplasm,  are  always  smaller  than  those  of  the  shell-valves  or 
of  polar  capsules.     Dimensions:  length  6  (?)  to  14jli,  breadth  4  to  6ju. 

MYXIDIUM  GLUTINOSUM  Davis 
[Fig.  291] 
1917        Myxidium  glutinosum  Davis  1917  :  235 

Habitat:  Gall-bladder  of  Cynoscion  regalis;  Beaufort. 


116  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [354 

Vegetative  form :  Elongated  or  irregular.  Slowly  ameboid,  moving  by- 
means  of  a  broad,  lobose  pseudopodium  of  hyaline  ectoplasm.  Body 
colorless.  Ectoplasm  only  distinct  in  pseudopodium.  Endoplasm  finely 
granular.  The  mature  spores  while  still  within  the  mother  trophozoites, 
are  surrounded  by  a  clear,  refractive  gelatinous  envelope.  Diameter  of 
rounded  sporulating  trophozoites  20/i.    Disporous. 

Spore:  Cylindrical,  ends  of  valves  rounded  except  at  one  side,  where 
the  polar  capsules  open  at  the  apex  of  a  small,  conical  elevation.  Spore 
characterized  by  the  presence  of  a  transparent,  homogeneous,  gelatinous 
envelope.  Polar  capsules  pyriform,  opening  on  each  side  nearly  at  right 
angles  to  the  longitudinal  axis.  Dimensions:  length  10  to  11/x,  breadth 
6/i,  length  of  polar  capsules  3/x. 

MYXIDIUM  PHYLLIUM  Davis 

[Figs.  292  and  293] 
1917        Myxidium  phylllum  Davis  1917  :  235 

Habitat:  Gall-bladder  of  Gamhusia  affinis;  Beaufort. 

Vegetative  form:  Exceptionally  large;  flattened,  leaflike,  usually  folded 
on  itself;  motionless.  Pseudopodia  were  not  observed.  Ectoplasm  forming 
a  distinct  transparent  layer  around  entire  body.  After  being  on  slide  for 
some  time  ectoplasm  usually  becomes  covered  with  very  numerous,  short, 
hairlike  processes.  Endoplasm  finely  granular,  contains  numerous  fat 
globules.    Diameter  up  to  1.35mm.    Polysporous. 

Spore:  Fusiform,  slightly  truncated  at  each  end  where  polar  capsules 
open.  Shell  with  numerous  longitudinal  striations.  Sporoplasm  finely 
granular,  with  several  small  fat  globules.  Dimensions:  length  1 1/x,  breadth 
8/x,  diameter  of  polar  capsules  3^. 

MYXIDIUM  STRIATUM  Cunha  et  Fonseca 
1917        Myxidium  striatus  Cunha  et  Fonseca  1917:321 

Habitat:  Gall-bladder  of  Menticirrhus  americanus  L.,  Bairdiella 
ronchus  Cuv.  et  Val. ;  Brazil. 

Vegetative  form:  More  or  less  spherical.  Body  small  and  colorless. 
Endoplasm  granular.  Ectoplasm  visible  when  pseudopodia  are  formed. 
Pseudopodia  filiform,  being  projected  radially.  Size  variable,  16ju  in 
diameter  in  average. 

Spore:  Elliptical.  Shell  with  fine  longitudinal  striations  which  run 
parallel  to  sutural  line.  Sutural  plane  oblique  to  the  longitudinal  axis  of 
the  spore  which  is  thickened  at  the  extremities.  Two  ovoidal  polar  cap- 
sules, one  at  each  end.  Dimensions:  length  10  to  14^t,  breadth  6  to  8/i, 
length  of  polar  capsules  4/i,  length  of  polar  filament  30/i. 


3551  STUDIES  ON  MYXOSPORIDIA—KUDO  117 

MYXIDIUM  KAGAYAMAI  nov.  spec. 
[Figs.  294  and  295] 
1916        Myxidium  sp.  Kudo  1916  : 6 

Habitat:  Gall-bladder  of  Misgurnus  anguillicaudatus  Cant.;  Tokio 
(September),  2%  of  the  fish  examined  infected. 

Vegetative  form:  Not  observed. 

Spore:  Fusiform;  one  valve  being  more  convex  than  the  other.  Suture 
line  straight.  Shell  with  fine  longitudinal  striations.  Dimensions  in  fixed 
preparations:  length  15  to  18m,  breadth  6  to  Ijx,  length  of  polar  capsules 
7  to  8m,  length  of  polar  filament  60  to  70^. 

Remarks:  Tho  the  vegetative  form  is  still  unobserved,  the  author  is 
compelled  to  consider  the  present  form  as  a  new  species  by  careful 
reexamination  of  the  material  and  proposes  the  name  in  honor  of  Dr. 
T.  Kagayama,  Tokio,  Nippon. 

MYXIDIUM  AMERICANUM  nov.  spec. 
[Figs.  622  to  627] 

Habitat:  In  the  lumen  of  urinary  tubules  of  the  kidney  of  Trionyx 
spinifera;  Crystal  Lake,  Urbana,  111.  (July).  A  single  host  specimen 
showed  a  light  infection  in  the  above  mentioned  organ.  No  intracellular 
stage  was  detected. 

Vegetative  form:  The  young  trophozoite  in  the  lumen  of  the  tubule 
of  the  kidney  is  multinucleate,  and  more  or  less  irregular  in  shape  which 
suggests  the  ameboid  movements  of  the  animal  (Figs.  622,  623).  The 
older  form  with  mature  spores  is  rather  spherical  in  form  with  a  distinct 
outline.  The  protoplasm  is  fairly  well  differentiated  into  ectoplasm  and 
endoplasm  (Fig.  624).  The  size  of  the  trophozoites  varies  from  12  to  25/1 
in  diameter.     A  pansporoblast  produces  two  spores.     Polysporous. 

Spore:  Spindle-form;  with  the  two  pointed  extremities  stretched  in 
opposite  directions.  Circular  in  cross-section.  The  shell  is  rather  thin; 
sutural  line  is  straight.  Fine  longitudinal  striations  on  the  shell,  eight 
to  ten  in  number  on  each  valve.  The  polar  capsules  are  nearly  spherical, 
coiled  polar  filament  being  visible  in  fresh  material  (three  turns).  The 
polar  filament  is  easily  extruded  from  the  fresh  spores  under  the  influence 
of  potassium  hydrate  solution.  The  direction  of  the  extruded  polar  fila- 
ment forms  an  angle  of  about  45°  with  the  main  axis  of  the  spore  and  the 
two  filaments  are  parallel  to  each  other.  Preserved  spores  do  not  show 
any  filament  extrusion  under  the  influence  of  the  said  chemical.  The 
sporoplasm  is  finely  granular,  and  shows,  upon  staining,  two  small  nuclei 
of  ring-shape,  as  their  peripheral  layer  takes  stain  more  deeply  than  the 
central  portion.  Average  dimensions  of  fresh  spores:  length  15  to  16m, 
breadth  5.5  to  6m,  polar  capsule  4m  by  3.5m,  length  of  polar  filament 
25  to  32m. 


118  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [356 

Remarks:  Two  species  of  the  genus  Myxidium  were  reported  to  occur 
in  chelonian  hosts;  i.e.,  M.  danilewskyi  (page  109)  and  M.  mackiei  (page 
112).  The  former  diflFers  from  the  present  form  in  having  an  elongated 
vegetative  form  which  is  greenish  in  color,  and  in  having  spores  of  different 
shape,  dimensions  and  structure,  not  to  speak  of  the  difference  of  the 
host.  The  latter  resembles  closely  to  the  species  under  consideration  in 
dimensions  of  the  spores,  but  differences  in  the  trophozoite  and  in  the 
structure  of  the  spore  do  not  allow  one  to  consider  two  forms  as  identical. 
The  species  is  therefore  treated  as  new. 

Genus     SPHAEROMYXA     Thelohan 
1892        Sphaerotnyxa  Th61ohan  1892a  :  1091-1093 

The  characters  of  the  genus  are  described  on  page  58. 
Type  species:  Sphaerotnyxa  balbianii  Thelohan. 

SPHAEROMYXA  BALBIANII  Thelohan 
[Figs.  296  to  307] 

1892        Sphaeromyxa  balbianii  Thelohan  1892a  :  1091-1093 

1895        Sphaeromyxa  balbianii  Th61ohan  1895  :  342 

1912        Sphaeromyxa  balbianii  Parisi  1912  :  288 

1916  Sphaeromyxa  balbianii  Georg^vitch  1916  :  92-93 

1917  Sphaeromyxa  balbianii  Davis  1917  :  235-236 

Habitat:  Gall-bladder  of  Motella  tricirrata  Bl.,  M.  maculata  Risso., 
Cepola  rubescens  L.,  Clupea  pilchardus,  Siphostoma  floridae,  S.  louisianae; 
Roscoff  (September),  Concarneau,  Marseille,  Banyuls,  Napoli  (Septem- 
ber), Beaufort  (June  to  August). 

Vegetative  form:  Flattened  leaf -like  or  disc-form,  reaching  3  to  4mm. 
in  diameter.  Often  forms  spherical  with  opaque  appearance.  The  proto- 
plasm is  distinctly  differentiated  into  endoplasm  and  ectoplasm.  Ecto- 
plasm forms  rounded  lobes  which  exhibit  slow  movements  and  show  a  clear 
radially  striated  structure  in  sections.  Endoplasm  reticular,  contains 
nuclei,  young  and  mature  spores  and  fat  globules.  Each  pansporoblast 
develops  two  spores.    Polysporous. 

Davis  mentions  that  the  largest  form  he  observed  was  900^  in  diameter. 

Georgevitch  recognized  a  large  number  of  small  trophozoites  which  were 
formed  by  repeated  plasmotomous  multiplication. 

Spore:  Fusiform,  with  truncate  ends.  Shell  longitudinally  striated. 
One  polar  capsule  at  each  end.  Polar  filament  is  wound  around  an  imagi- 
nary axis  perpendicular  to  the  longitudinal  axis  of  the  spore.  When 
extruded,  the  polar  filament  is  seen  as  a  short,  conical  and  hollow  thread- 
like structure.  Sporoplasm  finely  granular  with  two  nuclei.  Dimensions: 
length  15m,  width  5fx,  length  of  polar  filament  15/Lt. 


357]  STUDIES  ON  MYXOSPORIDIA—KUDO  119 

Parisi  gave  the  following  dimensions:  length  15  to  20ju,  width  5  to  6/i, 
polar  capsule  7/i  by  4.7/x,  length  of  polar  filament  25  to  30jii. 

Davis'  measurements  are  as  follows:  length  17  to  20/i,  breadth  5  to  6/i, 
length  of  polar  filament  20/i. 

Georgevitch  observed  young  spores  with  both  ends  tapering  into  a 
point.  Later  they  assume  the  typical  form  with  truncated  ends.  He  did 
not  recognize  the  striations  on  the  shell.  He  also  mentions  the  occurrence 
of  abnormal  spores,  such  as  elliptical,  spherical  forms,  etc.;  or  with  only 
one  polar  capsule. 

SPHAEROMYXA  IMMERSA  (Lutz)  Thelohan 
[Figs.  308  to  311] 

1889        Cystodiscus  immersus  Lutz  1889  :  84-88 

1895        Sphaeromyxa  immersa  Thelohan  1895  :  343 

1899        Cystodiscus  immersus  Liihe  1899  :  291-293 

Habitat:  Gall-bladder  of  Bufo  marinus  L.  and  Leptodactylus  ocellatus 
L.;  Brazil. 

Vegetative  form:  Leaf-like  or  disc  form,  visible  thru  the  bladder  wall. 
Upper  and  lower  sides  slightly  convex.  Size  up  to  1.5  or  2mm.  in  diameter; 
thickness  being  1/20  to  1/10  of  the  diameter.  Protoplasm  is  well  differ- 
entiated. Ectoplasm  transparent  and  membranous,  often  contains  a 
large  number  of  micrococcus-like  bodies.  No  ameboid  movements  nor 
change  of  form.  Endoplasm  highly  vacuolar,  contains  fat  globules. 
Plasmotomic  multiplication  probably  occurs.  Spores  always  arranged  in 
pairs.     Polysporous. 

Spore:  Oval  with  rounded  extremities.  Shell  more  or  less  thick,  with 
fine  transverse  striations.  Spherical  polar  capsule  at  each  end.  Sutural 
plane  is  oblique  to  the  longitudinal  axis  of  the  spore.  Sporoplasm  trans- 
parent. Dimensions:  length  12  to  14/x,  breadth  9  to  10/x,  length  of  polar 
filament  50  to  70/11  (4  to  5  times  that  of  the  spore)  (KOH). 

SPHAEROMYXA  INCURVATA  Doflein 

[Figs.  312  to  314] 

1898        Sphaeromyxa  incurvaia  Doflein  1898  :  286-287 

Habitat :  Gall-bladder  of  Blennius  ocellatus ;  Napoli. 

Vegetative  form:  Trophozoites  are  found  in  large  masses  (Plasmodia  ?), 
in  which  they  form  a  thin,  hollow  ball,  5-7mm.  in  diameter. 

As  the  surface  is  greater  than  the  inner  surface  of  the  bladder,  some 
parts  of  the  body  are  folded  up.  Body  bluish  white  and  transparent. 
Protoplasm  highly  vacuolar,  contains  numerous  fat  globules,  nuclei  and 
spores.     Polysporous. 

Spore :  Curved  to  one  side  along  sutural  plane  and  also  in  a  plane  at  right 
angles  to  it.     Polar  capsule  at  each  end.     Sporoplasm  with  two  nuclei. 


120  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [358 

Polar  filament  is  wound  along  the  longer  diameter  of  the  capsule,  and 
relatively  thick,  but  thinner  than  that  of  S.  balbianii  Thel.  Dimensions: 
length  (along  the  inner  side  of  the  arch)  30  to  35/*,  breadth  8/t,  distance 
between  two  polar  capsules  12  to  IS^u,  polar  capsules  12  to  15m  by  4  to  5/i. 


SPHAEROMYXA  SABRAZESI  Laveran  et  Mesnil 
[Figs.  315  and  322] 


1900 

Sphaeromyxa  sabrazesi 

Laveran  et 

Mesnil 

1900 

:  380-382 

1906 

Sphaeromyxa  sabrazesi 

Schroder 

1906  : 

:  455-466 

1907 

Sphaeromyxa  labrazesi* 

Schroder 

1907  : 

:  359-381 

1910 

Sphaeromyxa  sabrazesi 

Schroder 

1910; 

:l-5 

1912 

Sphaeromyxa  sabrazesi 

Parisi 

1912  ; 

:288 

1913 

Sphaeromyxa  sabrazesi 

Jameson 

1913  ; 

;2 

1916 

Sphaeromyxa  sabrazesi 

Georg6vitch 

1916; 

:  91-92 

1916 

Sphaeromyxa  sabrazesi 

Georg^vitch 

1916a 

:3 

Habitat:  Gall-bladder  of  Hippocampus  hrevirostris  Cuv.,  H.  guttulatus 
Cuv.;  Syngnathus  acus,  Motella  tricirrata,  Nerophis  annulatus,  Siphonos- 
toma  rondeletii;  Arcachon,  Rovigno,  Napoli,  Roscoff  (September),  Monaco, 
Villef ranee,  (March  to  June). 

Vegetative  form:  Disc  form.  Diameter  up  to  2mm.  Thickness  var- 
iable. Body  whitish  in  color.  Ectoplasm  thin,  transparent  and  homo- 
geneous. Young  trophozoites  may  probably  have  lobose  pseudopodia. 
Endoplasm  highly  vacuolated,  contains  nuclei  of  various  sizes,  pansporo- 
blasts, spores  and  more  or  less  refringent  granules.    Polysporous. 

Schroder  observed  larger  forms  up  to  5mm.  Ectoplasm  also  was  found 
to  project  numerous  fine  short  (Iju)  hair-like  processes  from  the  surface. 
Each  pansporoblast  develops  into  two  spores. 

Spore:  Cylindrical,  bent  in  arch  form;  with  truncated  ends.  Large 
cylindrical  polar  capsule  at  each  end.  Sporoplasm  granular,  contains  one 
nucleus.  Polar  filament  short  and  conical,  is  extruded  under  the  action  of 
nitric  acid.  Dimensions:  length  28)u,  width  4.3m,  polar  capsule  9  to  10m 
by  3n,  distance  between  the  polar  capsules  8m,  length  of  polar  filament  8m. 

Schroder  noticed  the  stained  sporoplasm  contained  one  or  two  nuclei. 
He  observed  indistinctly  marked  longitudinal  striations  on  the  shell. 
Dimensions:  length  22  to  25m,  breadth  3  to  4m,  polar  capsule  8m  by  2  to  3m, 
length  of  polar  filament  about  12m. 

Georgevitch  described  the  presence  of  a  hyaline  substance,  containing 
pale  granules,  in  the  spore  cavity.  Young  spores  were  found  to  take  the 
form  of  Myxidium  type.  In  mature  spores,  he  always  found  two  nuclei  by 
staining. 

*Misprmted  in  Schroder's  pap)er. 


359]  STUDIES  ON  MYXOSPORIDIA—KUDO  121 

SPHAEROMYXA  HELLANDI  Auerbach 
[Figs.  323  and  324] 

1909  Sphaeromyxa  hellandi  Auerbach  1909  :  78-79 

1910  Sphaeromyxa  hellandi  Auerbach  1910b  :  772-774 
1910        Sphaeromyxa  hellandi                Auerbach             1910c  :  174^175 

1912  Sphaeromyxa  hellandi  Auerbach  1912  : 4, 40 

Habitat:  Gall-bladder  of  Molva  vulgaris  Flem.,  Centronotus  gunellus, 
Brosmius  brosme  Asc;  Bergen,  Torghatten. 

Vegetative  form:  Large  and  rounded  disc  form.  Protoplasm  is  dis- 
tinctly differentiated  into  ectoplasm  and  endoplasm.  Thickness  up  to 
160/i,  folded  in  the  bladder.  Ectoplasm  finely  granular;  in  unstained  speci- 
mens, it  is  recognizable  as  10  to  12jli  thick  layer,  in  which  about  2n  thick 
radially  striated  outer  layer  and  8  to  10/x  thick  inner  finely  granular  region 
can  be  distinguished.  In  stained  sections,  the  outer  layer  remains 
unstained.  Endoplasm  highly  alveolar,  contains  refractive  granules  of 
different  size  which  are  not  stained  with  Sudan  III.  Each  pansporoblast 
develops  into  two  spores.    Polysporous. 

Spore:  Arch  form  in  front  view.  The  degree  of  curvature  varies 
greatly.  Sutural  line  curved  in  S-shape  and  well  marked.  Both  ends  more 
or  less  truncated.  Polar  capsule  at  each  end.  Polar  filament  being  wound 
along  the  longest  axis  of  the  polar  capsule  and  is  extruded  with  KOH. 
Sporoplasm  rounded  with  one  or  two  nuclei.  Dimensions:  length  20.8  to 
26;u,  breadth  and  thickness  5  An,  length  of  polar  capsule  10  to  lO.Sju. 

SPHAEROMYXA  EXNERI  Awerinzew 
[Figs.  325  and  326] 

1913  Sphaeromyxa  exneri  Awerinzew  1913a  :  155 

Habitat:  Gall-bladder  of  Thysanophris  japonicus;  Lorenzo  IVIarques 
(Africa). 

Vegetative  form:  Not  observed. 

Spore:  Somewhat  resembles  that  of  S.  hellandi  Auer.  in  being  bent 
to  one  side  with  sutural  line  of  S-form  but  differs  in  dimensions.  Both 
ends  slightly  tapering.  Polar  capsules  two,  one  at  each  blunt  end,  in 
which  the  polar  filament  is  wound  parallel  to  its  longer  axis.  Sporoplasm 
comparatively  small  and  sharp-contoured,  contains  only  one  nucleus. 
Dimensions:  length  75  to  80/x,  breadth  18  to  20/i,  length  of  polar  capsule 
30  to  35m. 

SPHAEROMYXA  GASTEROSTEI  Georgevitch 
[Fig.  327] 
1916        Sphaeromyxa  gasterostei  Georgevitch         1916  :  88 

Habitat:  Gall-bladder  of  Gasterosteus  spinachia;  Roscoff  (September). 
Vegetative  form:  Trophozoites  form  large  plasmodia. 


122  ILLINOIS  BIOLOGICAL  MONOGRAPHS  (360 

Spore:  Large,  elongated  fusiform;  ends  less  truncated  than  those  of 
the  spore  of  Sphaeromyxa  balbianii.  As  the  spore  becomes  mature,  the 
ends  assume  more  pointed  shapes.  Polar  capsules  two,  one  at  each  end. 
Sporoplasm  with  two  nuclei,  fills  the  extracapsular  cavity.  Dimensions: 
twice  or  three  times  larger  than  those  of  Sphaeromyxa  balbianii  Thelohan. 

Genus    ZSCHOKKELLA    Auerbach 
1910        Zschokkella  Auerbach  1910  :  62-63 

1910a  :  240-256 
1910c  :  175 

The  characters  of  the  genus  are  described  on  page  58. 
Type  species:  Zschokkella  hildae  Auerbach. 

ZSCHOKKELLA  HILDAE  Auerbach 
[Figs.  328  to  331] 
1910        Zschokkella  hildae  Auerbach  1910  :  62-63 

1910        Zschokkella  hildae  Auerbach  1910a  :  240-254 

1912        Zschokkella  hildae  Auerbach  1912  :  40-41 

Habitat:  Urinary  bladder  of  Phycis  blennioides  Br.,  Gadus  aeglefinis, 
G.  callarias  L.,  G.  virens  L.;  Norway. 

Vegetative  form:  Trophozoites  float  in  the  bile  or  attach  themselves 
to  the  epithelial  layer  of  the  bladder.  Youngest  ameboid  form  about 
4.5  to  6/x.  In  floating  form,  pseudopodia  more  or  less  long,  lobose,  are 
formed;  while  in  the  attached  form  those  similar  to  the  pseudopodia  of 
Myxidium  bergense  Auer.  are  developed.  Plasmogamy  occurs.  Size 
varies  greatly  according  to  the  number  of  spores  which  are  formed  in  each 
individual.  Monosporous  (with  or  without  the  remnant  of  protoplasm), 
disporous  and  polysporous  (up  to  4  spores). 

Spore:  Semicircular  in  front  view,  with  slightly  and  equally  attenuated 
ends.  At  each  end,  large  spherical  polar  capsule  is  situated  which  opens 
not  at  the  extremity,  but  on  the  flat  surface.  Shell  bivalve  and  thick. 
Sutural  line  S-form.  Sporoplasm  with  two  nuclei.  Dimensions:  length 
16  to  28.8/i,  breadth  13  to  18^i,  polar  capsules  5.6  to  7.2/i  in  diameter,  length 
of  polar  filament  72m  (KOH). 

ZSCHOKKELLA  NOVA  Klokacewa 
[Figs.  332  and  333] 
1914        Zschokkella  nova  Klokacewa  1914  :  184-186 

Habitat:  Gall-bladder  of  Carassius  vulgaris;  Russia? 
Vegetative  form:  Not  observed. 

Spore:  Outline  irregular.  Observations  on  fixed  materials  alone.  Two 
large  round  polar  capsules  open  at  the  side  near  ends.  Sporoplasm  with 
two  nuclei.  On  some  spores,  striations  that  run  parallel  to  the  sutural  line 
were  observed.  Dimensions:  length  9.5  to  11.5/i,  breadth  6.5  to  7/t, 
diameter  of  polar  capsule  3  to  3.5m. 


361]  STUDIES  ON  MYXOSPORIDIA—KUDO  123 

ZSCHOKKELLA  ACHEILOGNATHI  Kudo 
[Figs.  334  to  338] 

1916  Zschokkella  acheUognathi  Kudo  1916  : 3-5 

Habitat:  Gall-bladder  and  gall-duct  of  Acheilognathus  lanceolatum 
Temm.  et  Schl.;  Tokio  (May).  Over  80%  of  the  fish  examined  were 
found  to  be  infected. 

Vegetative  form:  Disc-shape.  In  bile  duct,  large  trophozoites  are 
folded  up.  Body  colorless  and  transparent.  Protoplasm  is  well  defined 
into  two  regions.  Ectoplasm  finely  granular  in  vivo.  In  stained  sections, 
it  shows  two  layers;  thin  outer  layer  (2/x  thick)  presents  very  fine  striations, 
while  inner  layer  (6  to  8/i  thick)  is  finely  vacuolated  without  any  enclosure. 
Endoplasm  is  highly  vacuolated.  Lobose  pseudopodia  formed  only  in 
younger  individuals  (15  to  30/i  in  diameter),  in  which  ameboid  movements 
are  not  slow.    Size:  up  to  720/i  by  550ju,  thickness  5  to  30/i.    Polysporous. 

Spore:  Form  resembles  Zschokkella  hildae,  but  slightly  elongated. 
Form  varies  to  some  extent.  Some  spores  are  of  Myxidium  type.  Sutural 
line  curved.  Longitudinally  striated.  Spherical  polar  capsule  at  each  end 
opening  near  the  extremity.  Dimensions:  length  10  to  14/*,  breadth  6  to  7/x, 
diameter  of  polar  capsule  3  to  5/x,  length  of  polar  filament  65  to  70/i  (KOH). 

ZSCHOKKELLA  GLOBULOSA  Davis 
[Figs.  339  and  340] 

1917  Zschokkella  globulosa  Davis  1917  :  236 

Habitat:  Urinary  bladder  of  Spheroides  maculalus;  Beaufort  (August). 

Vegetative  form:  Rounded;  slowly  ameboid,  forming  short,  lobose 
pseudopodia.  Body  colorless  and  transparent.  Ectoplasm  not  distinct. 
Protoplasm  granular,  characterized  by  the  presence  of  several  large  fat 
globules.  Sporulating  trophozoites  about  15  to  16/x  in  diameter.  Mono- 
sporous  and  disporous. 

Spore:  Semicircular.  Sutural  line  twisted  on  its  axis  and  oblique  to 
longitudinal  axis;  sutural  ridge  distinct.  Polar  capsules  opening  on  flat 
surface.  Sporoplasm  finely  granular  and  very  transparent.  Dimensions: 
length  11/x,  breadth  7/t,  diameter  of  polar  capsules  3/i. 

Family     MYXOSOMATIDAE     Poche 
1913        Myxosomatidae  Poche  1913  :  230 

The  characters  of  the  family  are  described  on  page  58. 

Genus    MYXOSOMA    Thelohan 

1892        Myxosoma  Th61ohan  1892  :  175 

The  characters  of  the  genus  are  described  on  page  58. 
Type  species:  Myxosoma  dujardini  Thelohan. 


124  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [362 

MYXOSOMA  DUJARDINI  Thelohan 


[Figs. 

341  to  343] 

1841 

MiiUer 

1841 

:486-487 

1845 

Dujardin 

1845 

:644 

1892 

Myxosoma  dujardini 

Thdohan 

1892 

:175 

1895 

Myxosoma  dujardini 

Thflohan 

1895 

-.343-344 

1905 

Myxosoma  dujardini 

Nufer 

1905 

:  77,  79, 186 

1910 

Myxosoma  dujardini 

Wegener 

1910 

:  72-73 

1916 

f  Myxosoma  dujardini 

Kudo 

1916 

:3 

Habitat:  Branchial  lamellae  of  Scardinius  erythrophthalmus  L.,  Perca 
fiuviatilis,  Leuciscus  rutilus  L.  and  Cyprinus  carpio  L.;  France,  Frisches 
Haff,  Kurisches  HafF  (February,  April,  May),  Tokio  (May),  Switzerland. 

Vegetative  form:  White  cysts  being  branched,  rounded,  spherical  or 
irregular;    1  to  1.5mm.  in  diameter. 

Wegener's  form  1  to  1.7mm.  long. 

Spore :  Ovoidal,  flattened,  with  attenuated  anterior  end  which  is  slightly 
bent  laterally.  Two  pyriform  polar  capsules  at  the  anterior  end.  Sporo- 
plasm  without  any  iodinophilous  vacuole.  Dimensions:  length  12  to 
13n,  breadth  7  to  8/i. 

Wegener's  form:  polar  capsules  6jj,  by  3/*. 

Kudo's  form:  polar  capsules  6  to  Ifx  by  2/i,  length  of  polar  filament  TO/*. 

MYXOSOMA  (?)  LOBATUM  Nemeczek 

[Fig.  348] 

1911        Myxosoma  (?)  lobalum  Nemeczek  1911  :  160-162 

Habitat:  Branchiae  of  Leuciscus  leuciscus  L.  and  Aspius  rapax  Ag.; 
Austria. 

Vegetative  form:  Cysts  spherical,  oval  or  elongated;  of  white  color. 
Size  from  0.5  to  3mm.  by  0.5  to  1mm.  Those  in  Aspius  rapax,  oval  to 
spindle-shape,  1  to  3mm.  long  and  1  to  1.5mm.  wide. 

Spore:  Ovoidal;  anterior  end  narrowly  pointed  and  straight;  posterior 
end  rounded,  with  lobose  appendix  (about  6/x  long).  A  transverse  fold 
on  the  shell  behind  the  polar  capsules  in  fresh  as  well  as  preserved  spores. 
No  iodinophilous  vacuole.  Dimensions:  length  12. 6/x,  breadth  8.2/t, 
length  of  polar  capsule  4.2^,  length  of  polar  filament  80  to  90/x.  Spores 
found  in  Aspius  rapax,  had  slight  difference  in  dimensions,  the  structure, 
however,  being  similar  to  the  above. 

Remarks:  Nemeczek  doubts  if  this  form  is  really  Myxosoma  because 
of  the  following:  1)  different  shape  of  the  cysts  compared  with  that  of 
the  type  species  as  described  by  Thelohan;  2)  spores  observed  might 
develop  later  into  other  forms  like  Henneguya. 


363]  STUDIES  ON  MYXOSPORJDIA— KUDO  125 

MYXOSOMA  FUNDULI  Kudo 
[Figs.  344  to  347] 
1918        Myxosoma  fundidi  Kudo  1918  :  12-14 

Habitat:  Branchiae  of  Fundulus  majalis  Wal.  and  F.  heteroclitus  L; 
Woods  Hole  (August,  September). 

Vegetative  form:  Cysts.  Spherical  and  small;  150/*  in  average  diame- 
ter. Largest  form  observed  360/x  by  264)u.  Spores,  young  and  mature, 
were  found  in  the  cysts.    Polysporous. 

Spore:  Pyriform.  Shell  uniformly  thick  with  7  to  10  folds  on  sutural 
edge  at  the  posterior  portion.  Sutural  ridge,  fairly  well  marked.  Two 
polar  capsules  pyriform  and  of  equal  size  at  the  anterior  end.  Sporoplasm 
finely  granular  with  two  nuclei  but  without  any  iodinophilous  vacuole. 
Dimensions:  length  14/*,  breadth  8/i,  thickness  6/i,  polar  capsule  8/i  by  2yLi, 
length  of  polar  filament  38  to  42/i  (perhydrol,  KOH). 

Remarks:  The  writer  could  not  find  any  evidence  of  an  iodinophilous 
vacuole  by  treatment  with  various  iodine  mixtures,  which  is  the  most 
important  characteristic  of  the  genus.  Hahn's  form  {Myxobolus  funduli, 
p.  151)  should  be  distinguished  from  the  present  form. 

Genus     LENTOSPORA     Plehn 
1905        Lentospora  Plehn  1905  :  150 

The  characters  of  the  genus  are  described  on  page  58. 
Type  species:  Lentospora  cerebralis  (Hofer)  Plehn. 

LENTOSPORA  CEREBRALIS  (Hofer)  Plehn 


[Figs.  349  to  354] 

1903 

Myxobolus  cerebralis 

Hofer 

1903  : 

8 

1904 

Myxobolus  chondrophagus 

Hofer 

1904: 

:53 

1905 

Lentospora  cerebralis 

Plehn 

1905  ; 

;  145-166 

1909 

Lentospora  cerebralis 

Plehn 

1909  : 

:38 

1910 

Lentospora  cerebralis 

Auerbach 

1910c 

:176 

Habitat:  Cartilage  and  perichondrium  of  Trulta  iridea  Gibb.,  Salmo 
fontinalis  Mitch.,  Trutta  salar  L.;  Germany  (Karlsruhe  and  other  localities). 

Vegetative  form:  Ameboid  form.  Size  varies  greatly.  Small  ameboid 
form  probably  grows  up  into  large  individual  which  has  often  fifty  or  more 
ringform  nuclei  and  breaks  up  into  numerous  small  forms  by  division.  No 
sporous  character  is  observed  except  a  figure  of  a  disporous  form. 

Spore:  Circular  in  front  view;  lenticular  in  side  view,  with  more  or  less 
extensive  variation  in  length  and  breadth.  Shell  smooth.  Sutural  ridge 
distinctly  thickened.  Two  polar  capsules  pyriform  and  convergent,  are 
usually  of  same  size.  Extruded  polar  filaments  cross  each  other.  Sporo- 
plasm with  two  ring-form  nuclei  but  without  any  iodinophilous  vacuole. 


126  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [364 

Dimensions:  diameter  6  to  10/*,  length  of  polar  capsule  2/5  that  of  the 
spore,  length  of  polar  filament  40  to  SOfi  (limewater,  1%  KOH). 

Remarks:  Plehn  noticed  that  the  present  form  causes  the  chronic 
form  of  "Drehkrankheit"  among  young  fish  in  German  waters.  She  was 
unable  to  extrude  the  polar  filament  with  mineral  (?)  acids.  Auerbach, 
however,  could  extrude  the  filament  by  means  of  acids. 

LENTOSPORA  MULTIPLICATA  Reuss 

[Fig.  355] 
1906        Lentospora  multiplicata  Reuss  1906  :  203 

Habitat:  Muscle  of  Idus  melanotus  Heck. ;  Volga?,  Russia. 

Vegetative  form:  Not  described. 

Spore:  Oval.  Sutural  edge  broad  with  many  folds.  No  iodinophilous 
vacuole.  Dimensions:  length  12^,  breadth  9.5m,  thickness  6/i,  polar 
capsules  4/t  by  2.25/*. 

LENTOSPORA  ENCEPHALINA    Mulsow 

1911  Lentospora  encephalina  Mulsow  1911  :  483-485 

Habitat:  In  the  blood  vessel  of  the  brain,  especially  of  the  mid-brain  of 
Cyprinus  carpio  L.;  Munich  (spring).  Blood  vessels  are  the  only  seat  of 
infection.  In  most  cases  many  individuals  lie  parallel  to  one  another.  The 
infection  occurs  frequently  and  heavily.  The  effect,  however,  is  undeter- 
mined. 

Vegetative  form:  Trophozoite  elongated,  worm-like  and  circular  in 
cross  section.  The  body  is  covered  with  a  pellicula.  The  protoplasm  is 
distinctly  differentiated  into  homogeneous  ectoplasm  layer  and  inner  endo- 
plasm.    In  the  latter  are  found  numerous  granules,  small  nuclei  and  spores. 

Spore :  Almost  circular  in  front  view ;  profile  ?  No  iodinophilous  vacuole 
is  found.  The  polar  filament  is  easily  extruded  by  means  of  a  highly  diluted 
KOH  solution.    Diameter:  5  to  5.5^. 

LENTOSPORA  ASYMMETRICA  Parisi 
[Figs.  356  to  359] 

1912  Lentospora  asymmetrica  Parisi  1912  :  292-293 

Habitat:  Connective  tissue  of  kidney  of  Crenilabrus  pavo  C.  et  V.; 
Napoli  (September). 

Vegetative  form:  One  trophozoite  found;  a  small,  rounded  form  with 
thin  and  hyaline  ectoplasm  which  could  be  distinguished  from  the  endo- 
plasm  with  coarse  yellowish  globules,  containing  two  spores.    Disporous? 

Spore:  Oval  from  the  front;  flattened  and  fusiform  in  profile.  Sutural 
edge  with  many  triangular  folds,  which  are  more  clearly  seen  in  material 


365]  STUDIES  ON  MYXOSPORIDIA—KUDO  127 

preserved  in  formalin  than  in  fresh  condition.  Two  polar  capsules  of  same 
size,  are  situated  asymmetrically,  opening  at  the  side  near  the  anterior 
end.  Sporoplasm  granular  and  with  two  nuclei,  but  without  any  iodino- 
philous  vacuole.  The  polar  filament  not  being  extruded  by  ordinary 
reagents,  probably  because  the  spores  were  not  full-grown.  Dimensions: 
length  10  to  ll)u,  breadth  6.5  to  7/i,  length  of  polar  capsules  Six. 

LENTOSPORA  ACUTA  (Fujita)  Kudo 

[Figs.  360  to  362] 

1912        Sphaerospora  acuta  Fujita  1912  :  260-261 

Habitat:  Epithelium  of  branchial  lamellae  of  Carassius  auratus  L.; 
Sapporo,  Nippon. 

Vegetative  form:  Fujita's  description  is  simply  as  follows:  Sporoblast 
contains  about  two  spores. 

Spore:  Spherical  in  front  view,  with  slightly  pointed  anterior  end; 
spindle  shaped  in  side-views.  Shell  thin  and  smooth.  Two  convergent 
polar  capsules  are  of  different  sizes,  occupying  about  5/8  in  space  of  the 
spore.  No  vacuole  could  be  made  out  in  sporoplasm.  Dimensions: 
length  8  to  lO^t,  breadth  7  to  Sn,  thickness  5  to  6/x,  polar  capsules  5^  by  Afi. 

Remarks:  This  species,  recorded  incompletely  by  Fujita  as  Sphaero- 
spora, shows  characters  of  the  genus  Lentospora  in  spore  form  so  that  it 
is  provisionally  placed  here. 

LENTOSPORA  DERMATOBIA  Ishii 
[Figs.  594  to  596] 
1916        Lentospora  dermatobia  Ishii  1916  :  472-474 

Habitat:  In  the  integument  of  Anguilla  japonica  Temm.  et  Schl.; 
Shizuoka,  Nippon.  From  the  same  specimen  which  harboured  Myxidium 
a«gwj^/ae,  see  page  114.  The  number  of  cysts  reaches  probably  "several 
hundreds." 

Vegetative  form:  Cysts,  beneath  the  epidermis,  usually  subcircular, 
more  or  less  irregularly  triangular  or  quadrilateral  under  the  magnifier, 
with  the  largest  diameter  of  from  142  to  267/*.  The  epidermis  is  slightly 
lifted  up  by  the  cyst.  No  chromatophore  on  the  surface  of  the  cyst.  The 
cysts  separated  from  each  other,  are  found  mostly  in  the  central  region  of 
the  body,  head  and  fins  being  free  from  cysts.  In  cross-section,  cysts 
exhibit  oval  or  lenticular  shape  with  the  longest  diameter,  which  is  twice 
as  long  as  the  depth,  placed  parallel  to  the  surface  of  the  skin.  No  partic- 
ular pathological  change  was  noticed. 

Spore:  Circular  in  front  view;  broad  fusiform  or  lenticular  in  side  view. 
Sutural  ridge  fairly  distinct.    Sutural  edge  comparatively  broad,  especially 


128  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [366 

at  the  posterior  margin,  where  a  few  folds  (three  are  figured)  are  seen.  Two 
oval  polar  capsules  convergent  and  of  equal  size.  Sporoplasm  is  sharply- 
contoured,  no  iodinophilous  vacuole  being  recognized.  Dimensions  in 
preserved  material  (?):  diameter  6.3  to  7n,  thickness  4.2  to  4.9/u,  length 
of  polar  capsule  2.8  to  3.5/x. 

Family    MYXOBOLIDAE     Thelohan 

1892  Myxobolidies  Thdlohan  1892  :  173,  176 

1893  Myxoholidae  Gurley  1893  :  412, 413 
1895        Myxobolidies                            Thdohan  1895  :  347 

The  characters  of  the  family  are  described  on  page  58. 

Genus     MYXOBOLUS     Biitschli 
1882        Myxobolus  ButschU  1882  :  PI.  38  : 6-10 

The  characters  of  the  genus  are  described  on  page  58. 
Tjrpe  species:  Myxobolus  miilleri  Biitschli. 

MYXOBOLUS  MULLERI  BiitschH 


[Figs. 

397  to  4031 

1881 

Batsrhli 

1881 

630 

1882 

Myxobolus  miilleri 

Butschli 

1882 

595 

1895 

Myxobolus  miilleri 

Thdlohan 

1895 

349 

1905 

Myxobolus  miiUeri 

Nufer 

1905' 

77,  79,  186 

1906 

Myxobolus  miiUeri 

C6p^de 

1906 

64-65 

1906 

Myxobolus  miiUeri 

Schroder 

1906 

195 

1908 

Myxobolus  miilleri 

Auerbach 

1908 

:456 

1909 

Myxobolus  miiUeri 

Auerbach 

1909a 

:54,  71 

1910 

Myxobolus  miiUeri 

Wegener 

1910 

:81 

1911 

Myxobolus  miiUeri 

Nemeczek 

1911 

160 

1912 

Myxobolus  miiUeri 

Parisi 

1912 

:293 

Habitat:  Air  bladder  and  branchiae  of  Leuciscus  cephalus  L.;  kidney 
and  ovary  of  L.  phoxinus  L.;  eye  of  Crenilabrus  melops  L.  and  Alburnus 
lucidus;  branchiae  of  As  pro  as  per  L.,  Barbus  vulgaris  Flem.,  Leuciscus  ruii- 
lus  L.,  Squalius  cephalus  L.,  S.  agassizii  Heckel,  Lota  vulgaris  L.,  Phoxinus 
loevis  Ag. ;  pseudobranchiae  of  Cottus  gobio  L. ;  intestine  of  Mugil  auratus 
Risso.;  France,  Germany  [Karlsruhe,  AUe  (October),  Pregel,  Frisches 
Haff],  Switzerland  (Neuchatel  Lake),  Italy  (Napoli,  September). 

Vegetative  form:  White  cysts  in  the  connective  tissue.  Form  elongated 
oval,  2  to  3mm.  in  diameter.  No  clear  differentiation  of  protoplasm  is 
observed  even  in  young  forms.  In  sections,  some  cysts  show  radiate 
striations  in  the  thick  granule-free  ectoplasm.  Endoplasm  filled  with  nuclei. 

Cepede  writes  as  follows :  Cysts  in  branchiae,  subspherical  or  elliptical, 
1.5mm.  by  0.5mm. 

Wegener's  form:  Cysts  small  and  rounded,  0.2  to  0.3mm.  in  diameter. 

Spore:  Ordinarily    spherical   or   subspherical.     Two   polar   capsules 


367]  STUDIES  ON  MYXOSPORJDIA—KUDO  129 

with  a  small  triangular  intercapsular  appendix.  Polar  capsules  pyriform 
and  of  same  size.    Sutural  edge  exhibits  folds  (7  to  9). 

Thelohan's  dimensions:  length  10  to  12)Lt,  breadth  9  to  ll/x,  length  of 
polar  capsule  5/i. 

Cepede  gave  the  following  dimensions  in  vivo:  length  lO/x,  breadth 
9/x,  thickness  6/t,  length  of  polar  capsule  5/ix. 

Wegener's  form.  Usually  oval,  often  almost  spherical.  Length  10  to 
11/x,  breadth  8  to  9/i,  diameter  of  spherical  form  9^,  polar  capsule  4  to 
5/i  by  2  to  2>n. 

MYXOBOLUS  PIRIFORMIS  Thelohan 


[Figs.  363  to  364] 

1852 

Remak 

1852  : 

:144 

1883 

Balbiani 

1883 

:  197-198 

1884 

Balbiani 

1884: 

:125 

1891 

Pfeiffer 

1891 

:132 

1892 

Myxobolus  piriformis 

Th61ohan 

1892  : 

:177 

1893 

Myxobolus  piriformis 

Gurley 

1893  : 

:414 

1894 

Myxobolus  piriformis 

Gurley 

1894  ; 

:211 

1895 

Myxobolus  piriformis 

Th61ohan 

1895 

:348 

1905 

Myxobolus  piriformis 

Nufer 

1905 

:  77,  186 

1910 

Myxobolus  piriformis 

Plehn 

1910 

:  22-27 

1910 

Myxobolus  piriformis 

Wegener 

1910 

:73 

Habitat:  Branchiae,  spleen,  kidney  of  Tinea  tinea  L.,  Cohitis  fos silts  L. 
and  subcutaneous  connective  tissue,  spleen,  liver,  connective  tissue  of  the 
intestine  of  Leuciscus  sp.;  France,  Germany  (Pregel),  Switzerland. 

Vegetative  form:  Small,  long  thread-like  cysts.  Color  white.  Poly- 
sporous. 

Wegener's  form:  average  size,  length  1mm.,  breadth  0.09  to  0.1mm 

Spore:  Elongated  oval;  flattened.  Anterior  end  highly  attenuated 
and  slightly  bent  to  one  side.  One  pyriform  polar  capsule  at  this  end. 
Dimensions:  length  16  to  18/i,  breadth  7  to  8/i,  length  of  polar  filament 
30/x. 

Wegener  gives  the  following  dimensions:  length  18/x,  breadth  7. 5/i, 
polar  capsule  7. 5/i  by  3. 5/i. 

MYXOBOLUS  UNICAPSULATUS  Gurley 
[Figs.  365  to  366] 

1841  Miiller  1841  :  487 

1893  Myxobolus  unicapsulatus  Gurley  1893  :  414 

1894  Myxobolus  unicapsulatus  Gurley  1894  :  210-211 

Habitat:  In  the  skin  of  Labeo  niloticus  For.;  Nile. 

Vegetative  form:  Cysts  very  small  pustules  in  the  skin  of  the  head. 


130  ILLINOIS  BIOLOGICAL  MONOGRAPHS  (368 

Spore:  Form  similar  to  Myxosoma  dujardini.  A  single  polar  capsule 
at  the  anterior  end,  obliquely  directed.  Dimensions:  length  0.0051'", 
breadth  0.0034'". 

MYXOBOLUS  FUHRMANNI  Auerbach 
[Fig.  367] 

1909  Myxobolus  fuhrmanni  Auerbach  1909  :  65-68 

1910  Myxobolus  fuhrmanni  Auerbach  1910c  :  178-179 

Habitat:  Connective  tissue  under  the  mucous  membrane  of  the  mouth 
of  Leuciscus  rutilus  L.;  Neuchatel  Lake. 

Vegetative  form:  Cysts,  of  pea-size,  surrounded  by  several  membra- 
nous layers  of  connective  tissue  with  a  few  nuclei.  Finely  granular  ecto- 
plasm forms  outer  layer.  Endoplasm  is  dense  and  contains  faintly  stained 
nuclei.  Pansporoblasts  and  spores  are  found  in  the  central  portion  of  the 
cyst.    Polysporous. 

Spore:  Elongated  pyriform,  with  attenuated  anterior  and  rounded 
posterior  ends.  Majority  with  a  single  polar  capsule;  spores  with  two 
polar  capsules  were  also  observed.  Shell  thick,  at  the  posterior  end.  4  to 
6  notch-like  markings  on  the  posterior  part  of  the  shell.  Sutural  ridge 
thickened  and  fairly  well  marked.  Coiled  polar  filament  visible  in  pre- 
served material.  The  opening  of  the  polar  capsule  is  either  at  the  anterior 
end  or  near  it.  Sporoplasm  with  two  nuclei  of  unequal  size  and  a  com- 
paratively large  iodinophilous  vacuole,  stained  brown  with  iodine  alcohol. 
Dimensions:  length  18  to  20fx,  breadth  about  8fx,  thickness  6fi,  length  of 
polar  capsule  9  to  lO^t. 

MYXOBOLUS  OCULI-LEUCISCI  Trojan 
[Fig.  368] 

1909        Myxoholus  oculi-leucisci  Trojan  1909  :  679-682 

Habitat:  Vitreous  body  of  the  eye  of  Leuciscus  rutilus  L.;  Prague 
(May?). 

Vegetative  form:  Two  cysts,  spherical  and  subspherical,  100  to  180/i  in 
diameter.  Ectoplasm  finely  granular.  Outer  portion  of  endoplasm  with 
small  nuclei,  then  larger  nuclei  each  surrounded  by  protoplasm,  while  the 
central  portion  contains  spores.    Polysporous. 

Spore:  Elongated  oval,  flattened  dorso-ventrally.  Posterior  margin 
rounded.  At  the  anterior  end,  a  single  polar  capsule  with  distinctly  visible 
coiled  polar  filament.  Shell  smooth  without  any  markings.  Sporoplasm 
with  one  nucleus,  usually  elongated  oval  (2.8/i  in  diameter)  and  one 
vacuole,  occupies  more  than  half  of  the  space  of  the  spore.  Dimen- 
sions: length  9  to  lOjit,  breadth  4.5  to  5.5/i,  thickness  3n,  polar  capsule 
5ju  by  2fi. 


369)  STUDIES  ON  MYXOSPORIDIA—KUDO  131 

MYXOBOLUS  TOYAMAI  Kudo 
[Figs.  369  to  370] 

1915  Myxoholus  toyamai  Kudo  1915  :  517-523 
1917        Myxoholus  toyamai                   Kudo  1917  :  163-170 

Habitat: — Connective  tissue  of  branchial  lamellae  of  Cyprinus  carpio 
L.;  Tokio  (July). 

Vegetative  form :  Cysts,  ovoidal  or  in  shape  of  calabash.  Small  form  67 
by  50/1,  shows  clear  differentiation  of  protoplasm.  Ectoplasm  radially 
striated,  often,  differentiates  fine  processes  (2  to  3/i  long).  Endoplasm 
coarsely  granular,  contains  nuclei  from  1  to  4/i  in  diameter.  Size  up  to  190/x 
in  greatest  diameter  in  sections.  Two  spores  are  formed  in  each  pansporo- 
blast.    Polysporous. 

Spore:  Pyriform,  with  attenuated  anterior  and  rounded  posterior  ends. 
No  bilateral  symmetry.  Lateral  sides  are  curved.  Calabash  shaped 
spores  often  occur.  Shell  without  any  marking,  thickened  at  the  anterior 
end.  Sutural  ridge  shows  sometimes  a  short  {l.Six  long)  tail-like  process  at 
the  posterior  tip.  A  single  pyriform  polar  capsule  at  the  anterior  end;  in 
stained  preparations,  a  small,  oblong  mass  of  protoplasm  is  seen  between 
the  polar  capsule  and  the  shell.  Coiled  polar  filament  distinct.  Sporo- 
plasm  with  two  nuclei  of  usually  same  size  and  a  relatively  large  iodino- 
philous  vacuole,  3)U  in  diameter.  Dimensions:  length  \Sn,  breadth  7  to  8ju, 
thickness  5  to  6/i,  polar  capsule  7  to  8/*  by  3  to  4/i,  length  of  polar  filament 
40  to  45m  (pressure,  perhydrol,  KOH). 

MYXOBOLUS  NOTATUS  Mavor 
[Figs.  371  to  372] 

1916  Myxoholus  notatus  Mavor  1916a  :  70-71 

Habitat:  Connective  tissue  of  the  voluntary  muscles  on  the  sides  or 
tail  of  Pimephales  notatus  Raf.;  Georgian  Bay,  Canada  (Summer). 

Vegetative  form:  Cysts  as  large  as  3mm.  in  diameter,  are  surrounded 
by  a  layer  of  columnar  epithelial  cells  (origin  and  significance?)  and  a  dense 
layer  of  connective  tissue.  Protoplasm  is  not  clearly  differentiated,  tho 
the  cyst  is  surrounded  by  an  area  devoid  of  nuclei.  In  the  outer  region  of 
endoplasm,  numerous  nuclei  each  with  a  caryosome,  are  recognized.  In 
the  course  of  spore  formation,  two  nuclei  for  polar  capsules  appear  at  first, 
one  of  which  degenerates  later.    Polysporous. 

Spore:  Pyriform,  with  a  posterior  extension  forming  a  process,  5/i  in 
length  and  as  broad  as  the  spore.  A  single  polar  capsule  at  the  anterior 
end.  An  iodinophilous  vacuole  in  the  sporoplasm.  Dimensions:  length 
17  to  18/c,  breadth  7.5  to  8/t,  polar  capsule  7/t  by  4/i,  length  of  polar  filament 
95/i. 


132  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [370 

MYXOBOLUS  sp.  Kudo 
1918        Myxobolus  sp.  Kudo  1918  :  15 

Habitat:  Spleen  of  Perca  flavescens;  West  Falmouth,  Mass.  (August). 
Isolated  spores  were  noticed  in  one  fish,  in  smears  and  section  preparations. 

Vegetative  form:  Not  observed. 

Spore:  Ovoidal,  attenuated  at  the  anterior  end.  Shell  uniformly  thick. 
A  single  polar  capsule  opens  at  the  anterior  tip.  Sporoplasm  contains  an 
iodinophilous  vacuole  and  two  nuclei  of  equal  size  (2)u).  Dimensions: 
length  18  to  20/1,  breadth  8/x,  polar  capsule  7  to  9/*  by  3  to  6/x. 

MYXOBOLUS  ROHITAE  Southwell  et  Prashad 

[Figs.  373  to  374] 

1918        Myxobolus  rohitae         Southwell  et  Prashad  1918  :  344-347 

Habitat:  Branchiae  of  Labeo  rohita;  Turag  river,  Mirpur,  Dacca 
district,  Bengal  (June).  Type  specimens  of  Indian  Museum  P48/1. 
Infection  was  heavy.  In  one  case  SZ  cysts  were  found  on  one  surface  of 
a  single  gill. 

Vegetative  form:  Cysts  in  the  gill-filaments.  The  cysts  preserved  in 
alcohol  are  of  a  creamy-yellow  color,  oval  to  cylindrical  in  form,  lying  with 
the  long  axis  parallel  to  the  gill- filaments.  Cysts  attached  to  the  gill- 
filaments  with  the  flattened  surface.  Size:  length  3.1  to  3.8mm.;  breadth 
0.8  to  1.2mm.  Cyst- wall  striated  vertically,  covered  with  an  epithelium, 
two  to  three  layers  thick.  In  the  central  portion  and  at  the  periphery, 
mature  spores  and  pansporoblasts  as  well  as  immature  spores  were  found 
respectively.     Polysporous. 

Spore:  Elongated  pyriform,  rounded  at  the  posterior  end  and  acutely 
pointed  anteriorly.  Sutural  ridge  slightly  raised.  One  polar  capsule 
present,  being  of  conspicuous  size.  Coiled  polar  filament  is  distinctly 
observed  in  the  polar  capsule.  An  iodinophilous  vacuole,  3.6/x  in  diameter, 
in  the  sporoplasm.  "Lying  just  posterior  to  it  is  the  nucleus  of  the  spore. 
A  few  granules  of  chromatin  were  also  seen  lying  scattered  in  the  proto- 
plasm." Dimensions:  length  30  to  32/i,  breadth  7  to  8/x,  length  of  the  polar 
capsule  22  to  23/*,  that  of  polar  filament  92  to  97/i. 

MYXOBOLUS  SENI  Southwell  et  Prashad 

[Figs.  375  to  376] 

1918        Myxobolus  seni  Southwell  et  Prashad  1918  :  347 

Habitat:  On  the  median  and  caudal  fins  of  Labeo  rohita;  Mirpur, 
Dacca   (January).     Type  specimens  in  Indian  Museum  numbered  P  53/1. 

Vegetative  form:  Trophozoites  form  cysts  which  are  elongated  ellip- 
soidal.    Size  from  4.7mm.  to  5.4mm.  in  length,  2.9mm.  to  3.7mm.  in 


371] 


STUDIES  ON  MYXOSPORJDIA—KUDO 


133 


breadth.    Color  of  the  cyst  whitish  with  black  scattered  granules  on  the 
surface. 

Spore:  Oval,  much  wider  behind  than  in  front  and  pointed  at  the 
anterior  end.  Sutural  ridge  is  slightly  thickened.  A  single  polar  capsule, 
showing  much  coiled  polar  filament.  lodinophilous  vacuole  2.3m  in  diame- 
ter. Dimensions:  length  13.2  to  13.6/i,  breadth  10.1  to  10.3)li,  length  of 
polar  capsule  4/i,  length  of  polar  filament  43^1  (in  one  case) . 

MYXOBOLUS  MISGURNI  nov.  spec. 

[Figs.  377  to  378] 
1916        Myxobolus  fuhrmanni  Kudo  1916  :  S 

Habitat:  GaW-hladder  oi  Misgurnus  anguillicaudatus ;  Tokio  (Septem- 
ber). About  50%  of  the  fish  examined  showed  a  few  isolated  spores 
floating  in  the  bile. 

Vegetative  form:  Unobserved. 

Spore:  Form  elongated  pyriform,  with  attenuated  anterior  and  rounded 
posterior  ends.  Shell  uniformly  thick.  Over  sutural  edge,  shell  exhibits 
many  (up  to  12)  triangular  markings.  Sutural  ridge  distinct.  A  single 
pyriform  polar  capsule  at  the  anterior  end.  Sporoplasm  contains  an 
iodinophilous  vacuole  and  two  nuclei.  Coiled  polar  filament  distinct  in 
vivo.  Dimensions  of  fresh  spores:  length  14  to  15.5/x,  breadth  6  to  7.3^, 
thickness  5  to  6fi,  polar  capsule  6.3/i  by  2  to  3n,  length  of  polar  filament  up 
to  lOOju. 

Remarks :  The  writer  reported  this  species  as  identical  with  Myxobolus 
fuhrmanni  Auerbach.  By  repeated  reexamination  and  comparison  with 
Auerbach's  description,  however,  he  came  to  the  conclusion  that  the 
present  form  should  be  treated  as  a  new  species,  on  account  of  the  difference 
of  the  host  and  the  characters  of  the  spore. 


MYXOBOLUS  PFEIFFERI  Thelohan 
[Figs.  379  to  385] 


1890 

Myxosporidian 

Pfeiffer 

1890; 

: 30-37 

1891 

Myxosporidian 

Pfeiflfer 

1891  ; 

:  100, 105-110, 130 

1893 

Myxosporidian 

PfeiflFer 

1893; 

:  118-130 

1895 

Myxobolus  pfeifferi 

Th61ohan 

1895 

:350 

1898 

Myxobolus  pfeijfferi 

Doflein 

1898 

:  306,  320,  etc. 

1906 

Myxobolus  pfeijfferi 

C6pede 

1906 

:59 

1906 

Myxobolus  pfdferi 

Stazzi 

1906 

:14hl9 

1906 

Myxobolus  pfdferi 

Mercier 

1906 
1906a 

:  427^28; 
:  763-764 

1908 

Myxobolus  pfeiferi 

Keysselitz 

1908 

:  253-273, 
286-306 

1909 

Myxobolus  pfeiferi 

Mercier 

1909 

:.5-30 

134  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [372 

Habitat:  Muscle  and  connective  tissue  of  kidney,  spleen,  intestine, 
ovary,  etc.,  of  Barbus  barbus  L.,  and  branchiae  of  B.  fluviatilis  Ag.  and 
B.  plebejus  Val.;  Drac  (June),  Neckar,  Prag,  Milano.  The  cause  of  well 
known  ''Boil  disease"  (Beulenkrankheit)  or  Myxoboliasis  tuberosa  (Hofer) 
of  the  barbels  in  European  waters.  Among  many  observers  Keysselitz 
made  a  thoro  study  of  the  parasite.  His  observations  are  as  follows:  The 
disease  occurs  among  the  fish  at  any  stage  of  growth.  About  8%  of  the 
fish,  7  to  15cm,  long,  caught  in  May  and  June  between  Conz  and  Trier 
were  infected  with  the  parasites.  The  heaviest  infection,  however,  occurs 
among  fish  up  to  40cm.  in  length;  fish  50cm,  long  or  larger  show  the  tumors 
caused  by  the  parasites,  rather  rarely.  Most  of  the  fish  die  as  the  result 
of  the  infection  between  the  early  part  of  April  and  the  end  of  October. 
The  highest  mortality  is  reached  in  the  hottest  months,  i.e.,  July  and 
August.  The  temperature  greatly  affects  the  growth  of  the  parasites. 
Fish  kept  in  the  aquarium  at  a  temperature  of  25°  C.  or  higher  demon- 
strate the  growth  of  the  boil  in  size  daily.  The  boils  are  not  noticed 
during  the  winter  and  spring,  they  are  formed  from  the  early  part  of  April 
to  the  middle  of  October. 

Vegetative  form:  The  parasites  develop  tumors  of  conspicuous  size. 

Keysseltiz's  observations  are  as  follows:  The  tumor  varies  in  size  from 
millet-grains  to  hen's  eggs.  Form  spherical,  oval  or  elongated.  The 
number  of  cysts  on  a  single  fish,  is  usually  3  to  4;  often  one,  in  some  fish, 
however,  23  were  recognized  on  one  fish.  Usually  tumors  separated  from 
each  other,  rarely  many  forming  one  tumor.  In  one  fish,  27cm.  in  length, 
a  tumor  of  7cm.  long,  4cm.  broad  and  3cm.  thick,  was  observed  in  July. 
The  seat  of  infection  is:  the  muscle  of  the  body,  muscle  of  pectoral  and 
anal  fins,  often  in  peritoneum  and  rarely  in  intestine.  As  the  result  of 
breaking  up  of  the  cyst  membrane,  spores  are  also  found  in  the  testis,  liver 
and  kidney. 

The  tumor  is  composed  of  many  vegetative  forms,  rounded,  oval, 
elongated,  variously  branched  or  flattened.  Size  reaches  to  1.5mm.  in 
diameter.  Protoplasm  is  usually  differentiated  into  ectoplasm  and  endo- 
plasm.  The  surface  is  not  often  smooth,  but  shows  irregular  outline. 
Ectoplasm  is  seen  often  as  a  very  thin,  uniformly  hyaline,  indistinctly 
granular  or  radially  striated  layer,  giving  the  network-like  appearance  to 
the  surface  of  the  body.  Endoplasm,  stained  more  deeply  around  the 
peripheral  part  than  other  portion,  shows  a  coarsely  alveolar  structure  in 
the  central  region.  It  contains  vegetative  nuclei,  developmental  stages  of 
propagative  nuclei,  granules,  fat-like,  often  leucocytes  and  red  blood 
'corpuscles.  The  leucocytes,  uninuclear  or  multinuclear,  were  seen  at  the 
periphery,  apparently  in  the  course  of  degeneration.  Red  blood  corpuscles 
were  found,  in  section,  inside  of  the  apparently  intact  parasite.  Each 
pansporoblast  develops  into  two  spores.    Polysporous. 


373]  STUDIES  ON  MYXOSPORIDAJ—KUDO  135 

Cepede  observed  one  cyst,  about  2mm.  in  diameter,  in  the  connective 
tissue  of  the  third  gill  arch. 

Spore:  Thelohan  described  as  follows:  Ovoidal.  Sutural  edge  shows 
folds.  A  small  triangular  intercapsular  appendix.  Dimensions:  length 
12/x,  breadth  10/i.     Cepede's  form  showed  exactly  the  same  dimensions. 

Keysselitz  gave  the  characters  of  the  spore  as  follows: 

Flattened  oval.  Shell  smooth.  A  small  intercapsular  appendix. 
Sutural  edge  having  a  number  of  small  flat  enlargement,  size  and  number 
being  variable.  Two  convergent  narrow  canals  (foramina)  penetrate  the 
shell  at  the  anterior  end.  Two  polar  capsules,  pyriform  and  of  equal  or 
nearly  equal  size,  are  located  at  the  anterior  half.  Coiled  polar  filament 
distinct,  coiled  7  to  8  times.  No  distinct  connection  between  polar 
capsule  and  the  filament.  Sporoplasm  fills  the  posterior  half  of  the  spore, 
extending  into  intercapsular  cavity.  It  is  finely  reticular,  exhibits  one  or 
two  rounded  or  oval  vesicular  nuclei  and  an  iodinophilous  vacuole.  Fat- 
like substance  is  often  seen  around  the  polar  capsules.  Spores  kept  in  water 
for  four  months  remain  intact  in  large  numbers.  Dimensions:  length  12 
to  12.5/x,  breadth  10  to  lO.S/it,  length  of  polar  capsule,  5.5  to  6/x,  length 
of  polar  filament  28  to  34)u. 

MYXOBOLUS  INAEQUALIS  Gurley 
[Fig.  411] 

1841  MUUer  1841  :  487-488 

1893  Myxobolus  inaequalis  Gurley  1893  :  414 

1894  Myxobolus  inaequalis  Gurley  1894  :  212 

Habitat:  In  the  skin  of  the  head  of  Piramutana  blochi  Cuv.  et  Vil.  and 
Synodontis  schall  Bl.  Schn.;  Guiana,  Surinam. 

Vegetative  form:  Very  small  pustules  in  the  skin  of  the  head. 

Spore:  Ovoidal.  Two  polar  capsules  of  unequal  size  at  the  anterior 
end.    Dimensions:  length  0.0052'",  breadth  0.0033'". 


MYXOBOLUS  DISPAR  Thelohan 

[Fig.  386] 

1895 

Myxobolus  dispar 

Thelohan 

1895 

:348 

1904 

Myxobolus  dispar 

Hofer 

1904  ; 

:50 

1910 

Myxobolus  dispar 

Wegener 

1910 

:  73-74 

1911 

Myxobolus  dispar 

Nemeczek 

1911 

:145 

Habitat:  Branchiae  of  Carassius  carassius  L.,  branchiae  and  epithelium 
of  intestine  of  Cyprinus  carpio  L.,  also  muscle  and  spleen  of  Scardinius 
erythrophthalmus  L.  and  in  the  skin  and  the  connective  tissue  of  Alhurnus 
lucidus  Heck.;  France,  Austria,  Konigsburg  (March,  July,  September). 

Vegetative  form:  Not  described  by  Thelohan. 

Wegener's  description  is  as  follows:  Cysts:  white  in  color;  spindle  shape 
with  pointed  ends.     Cysts  in  Carassius  carassius  L.  smaller  and  oval. 


136  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [374 

Size  3.5mm.  by  0.8mm.  Cysts  are  surrounded  by  thick  layers  (7  to  8/li)  of 
the  connective  tissue  of  the  host.  Ectoplasm  seems  to  be  undifferentiated. 
Endoplasm  granular,  contains  a  larger  number  of  spores.    Polysporous. 

Spore:  Thelohan's  diagnosis  is  as  follows: 

Ellipsoidal  or  slightly  oval.  Shell  with  3  to  5  folds  along  sutural  edge. 
Polar  capsules  of  unequal  size,  mth  a  small  intercapsular  body.  The 
vacuole  is  difficult  to  stain  with  iodine.  Dimensions:  length  10  po 
12/i,  breadth  Sju,  polar  capsule  Tfx  by  5/x.  • 

Wegener's  form  is  as  follows:  length  11  to  12/i,  breadth  7.5  to  8/x, 
larger  polar  capsule  6  to  7n  by  3.5jLt,  smaller  one  4yn  by  2.5  to  3/i.  The 
sporoplasm  is  shifted  toward  the  smaller  polar  capsule. 

MYXOBOLUS  ELLIPSOIDES  Thelohan 


[Figs. 

387  to  389] 

1852 

Remak 

1852 

:  144-146 

1892 

Myxobolus  ellipsoides 

Th61ohan 

1892 

:177 

1895 

Myxobolus  ellipsoides 

Th61ohan 

1895 

:  350-351 

1898 

Myxobolus  ellipsoides 

Doflein 

1898  ; 

:  324,  etc. 

1905 

Myxobolus  ellipsoides 

Nufer 

1905 

:  77,  79,  186. 

1910 

Myxobolus  ellipsoides 

Wegener 

1910 

:74r-75 

1912 

Myxobolus  ellipsoides 

Lo  Giudice 

1912 

:l-79 

Habitat:  Connective  tissue  of  air  bladder,  branchiae,  kidney,  spleen, 
liver  and  cornea  of  Tinea  tinea  L.,  branchiae  oi  Ahramis  brama  L.,  Alburnus 
lucidus  Heck.,  Leueiseus  rutilus  L.,  Squalius  eephalus  L.,  Ahramis  vimpa 
Cuv.,  Blieea  bjorkna  L.,  Idus  melanotus;  France,  Vierwaldstatter  See, 
Prague,  Masurische  See,  Italy. 

Vegetative  form:  Thelohan  does  not  describe. 

According  to  Wegener,  white  cysts,  elongated  oval;  2mm.  by  0.5mm. 
in  size.    Polysporous. 

Spore:  Thelohan  described  as  follows:  Flattened  elliptical,  rather 
elongated.  Sutural  edge  broad  without  any  folds.  Shell  with  no  marking. 
Form  of  the  spore  somewhat  variable.  Two  polar  capsules  of  equal  size, 
capsulogeneous  nuclei  present  even  when  fully  grown.  Abnormal  spores 
are  of  frequent  occurrence.  Dimensions:  length  12  to  lAfi,  breadth  9  to 
ll/i,  length  of  polar  capsule  4/x. 

Wegener's  form:  length  14  to  15/i,  breadth  10  to  ll/t,  polar  capsule 
4  to  5n  by  3fi.    Shell  comparatively  thick.    One  spore  had  a  tail  5/i  long. 

MYXOBOLUS  EXIGUUS  Thelohan 


[Figs. 

390  to  395] 

1891 

Myxosporidium  mugilis  ? 

Perugia 

1891 

:23 

1895 

Myxobolus  exiguus 

Thaohan 

1895 

:  349-350 

1906 

Myxobolus  exiguus 

Schroder 

1906 

:195 

1910 

Myxobolus  exiguus 

Wegener 

1910 

:75 

1912 

Myxobolus  exiguus 

Parisi 

1912 

:  294-295 

375]  .       STUDIES  ON  MYXOSPORJDIA— KUDO  137 

Habitat:  Branchiae  of  Ahramis  brama  L.  and  Chondrostoma  nasus  L., 
wall  of  stomach,  pyloric  coecum  and  intestine,  branchiae,  spleen,  kidney  of 
Mugil  chelo  Cuv.,  M.  capita  Cuv.  and  M.  auratus  Riss.;  Le  Vivier-sur-mer, 
Banyuls,  Marseille,  Heidelberg,  Pregel,  Frisches  Haff,  Kurisches  HafiF, 
Geneva,  Napoli. 

Vegetative  form:  No  description  by  Thelohan. 

Wegener  writes  as  follows: 

Cysts  of  variable  size.  Color  white.  Usually  small  and  narrow,  0.5 
to  0.7mm.  long  and  0.2mm.  wide.  Frequently  large  round  cysts  of  1.2  to 
1.5mm.  in  diameter,  filling  the  lamella.  Cysts  are  surrounded  by  10  to 
1  l/i  thick  membrane  composed  of  the  connective  tissue  of  the  host.  Ecto- 
plasm, 5/x  thick,  is  faintly  stained  by  hematoxylin.  Outer  region  of  endo- 
plasm,  alveolar  and  densely  loaded  with  nuclei,  while  in  the  central  portion 
with  mature  spores  in  granular  ground-mass. 

Parisi's  observations  are  as  follows: 

Cysts  in  the  intestinal  wall  of  Mugil  auratus,  large;  reaching  a  length 
of  3mm. 

Spore:  Thelohan's  description  is  as  follows: 

Flattened  ovoidal,  with  more  or  less  attenuated  anterior  end.  Sutural 
edge  shows  fairly  noticeable  folds.  A  small  triangular  intercapsular  appen- 
dix. Vacuole  in  the  sporoplasm  is  usually  hard  to  stain  with  iodine. 
Dimensions:  length  8  to  9^i,  breadth  6  to  In,  length  of  polar  filament 
15/i  (KOH). 

Wegener  observed  as  follows:  Rounded  with  slightly  pointed  anterior 
end.  Length  8  to  9. 5/x,  breadth  6  to  7.5ju,  polar  capsule  4.5/i  by  2  to  3/i. 
Shell  exhibits  small  folds  around  the  sporoplasm.  An  intercapsular  triangu- 
lar body  indistinctly  visible. 

Parisi  gave  the  following  dimensions:  length  8  to  8.5ju,  breadth  6  to  7/i, 
thickness  5.5m,  polar  capsule  3  to  An  by  1.5  to  2ju,  length  of  polar  filament 
30ybt  (alkaline).     Folds  usually  6  in  number.    Coiled  polar  filament  visible 


m  vivo. 

MYXOBOLUS  OVIFORMIS  Thelohan 

[Fig.  396] 

1854 

Lieberkahn          1854 

:  21-22 

1892 

Myxobolus  oviformis 

Th61ohan             1892 

:177 

1895 

Myxobolus  oviformis 

Th6Iohan             1895 

:351 

1905 

Myocobolus  oviformis 

Nufer                   1905  ; 

:  77,  186 

1906 

Myxobolus  oviformis 

CdpMe                1906 

:60 

1910 

Myxobolus  oviformis 

Wegener              1910 

:  76-78 

Habitat:  Fin  (subcutaneous  tissue),  spleen,  kidney  and  liver  of  Gobio 
gobio  L.;  branchiae  of  Alburnus  lucidus  Heck.,  Cyprinus  carpio  L.,  Blicca 
bjorkna  L.,  Abramis  brama  L.  and  A.  vimba  L. ;  France  (Isere),  Frisches 
HafiF  (especially  spring  months),  Switzerland. 


138  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [376 

Vegetative  form:  Thelohan  gave  no  description. 

Wegener's  observations  are  as  follows: 

Cysts,  white,  0.75  to  1.7mm.  by  0.4  to  0.7mm.  In  sections,  cysts  are 
shown  to  be  surrounded  by  a  thick  (10  to  20/*,  average  16/*)  layer  of  connec- 
tive tissue.  Ectoplasm  a  thin  (6  to  8/i  thick)  layer,  exhibits  a  transverse 
striation.  The  striation  is  often  absent  at  places  in  ripe  cysts.  Endoplasm 
finely  granular.  In  young  cysts,  it  is,  however,  reticulated,  with  nuclei  of 
1.5/i  in  diameter. 

Spore:  Thelohan  described  as  follows:  Flattened  ovoidal  with  pointed 
anterior  end.  Shell  smooth.  No  folds.  Polar  capsule  comparatively  large. 
Dimensions:  length  10  to  12/i,  breadth  9/x,  polar  capsule  6/i. 

Cepede  observed  numerous  spores  in  the  liver  and  kidney  of  Gobio 
gohio.  Dimensions  in  vivo:  length  10  to  12/i,  breadth  9/*,  length  of  polar 
capsule  6/1.    Polar  capsules  of  equal  size.    Coiled  polar  filament  distinct. 

Wegener's  form:  length  10.5  to  ll/z,  breadth  7.5  to  8/i,  polar  capsule  5 
to  6/i  by  Zn. 

Remarks:  Wegener  recognized  another  form,  which  seems  to  be  of  very 
rare  occurrence  and  which  can  not  be  distinguished  distinctly  from  the 
above  described  form.  Cysts  at  the  end  of  the  branchial  lamellae.  Size 
1.7  to  2mm.  in  largest  length.  Spore  resembles  more  closely  the  figure 
given  by  Thelohan  for  Myxobolus  oviformis  than  the  above  mentioned  form 
which  he  observed.  A  small  intercapsular  appendix  (rounded)  indistinct. 
Sporoplasm  comparatively  small.  Length  12.5  to  13.5/*,  breadth  9/*, 
polar  capsule  7.5/i  by  3/*. 


MYXOBOLUS  LINTONI 

Gurley 

[Figs.  404  to  408] 

1891 
1893 

1894 

Linton 

Myxobolus  lintoni                     Gurley 
Myxobolus  lintoni                     Gurley 

1891 
1893 
1894 

:  99-102 

:414 

:238 

Habitat:  Superficial  musculature  and  subcutaneous  tissue  of  Cyprino- 
don  variegatus;  Woods  Hole  (August). 

Vegetative  form:  Cysts,  not  closed,  but  fungoid  masses  of  an  irregular 
shape,  varying  in  size  from  4mm.  by  2.5mm.  to  10mm.  by  4mm.,  projecting 
as  much  as  3mm.  above  general  surface  of  skin.  The  skin  of  the  host  overly- 
ing these  tumors,  is  more  or  less  cracked  and  broken,  the  scales  being 
scattered. 

Spore:  Elliptical  in  the  front  view;  lenticular  in  side  view.  Shell 
thick.  Sutural  ridge  marked.  Two  polar  capsules,  convergent,  at  the 
anterior  end.  Spores  kept  in  alcohol,  extruded  polar  filaments  under  the 
action  of  iodine  water  and  sulphuric  acid.  Sporoplasm  with  a  large  iodino- 
philous  vacuole.    Dimensions:  length  13.9/i,  breadth  ll/x,  thickness  8/x. 


377)  STUDIES  ON  MYXOSPORIDIA—KUDO  139 

MYXOBOLUS  GLOBOSUS  Gurley 
[Figs.  409  and  410] 

1893  Myxoholus  globosus  Gurley  1893  :  415 

1894  Myxobolus  globosus  Gixrley  1894  :  241 

Habitat:  Branchial  lamellae  of  Erimyzon  sucetta  ohlongus  Lac.  (Catos- 
tomus  tuberculatus  Le  Sueur);  Kinston  (N.C.),  Columbia,  (S.C.,  March), 
tributaries  of  Fox  River. 

Vegetative  form:  Cysts,  whitish,  elongated  elliptical  or  rod-shaped, 
surrounded  by  very  thin  membrane?  Size  up  to  0.5mm.  in  max.  length. 
PolysporoUs. 

Spore:  Globose,  subcircular  in  outline.  Shell  thin  and  very  transparent. 
Sutural  ridge  very  wide,  being  one  third  of  the  thickness  of  the  spore. 
Polar  capsules  two,  of  equal  size,  divergent.  Vacuole  present,  but  not 
clearly  contoured.    Dimensions:  length  7  to  8jw,  breadth  6  to  7/t,  thickness 

MYXOBOLUS  OBLONGUS  Gurley 

[Figs.  412  to  416] 

1841  Miiller  1841  :  487-490 

1893  Myxobolus  oblongus  Gurley  1893  :  414 

1894  Myxobolus  oblongus  Gurley  1894  :  234^38 

Habitat:  Beneath  the  skin,  chiefly  of  the  head  of  Erimyzon  sucetta 
ohlongus  Lac.  {Catostomus  tuberculatus  Le  Sueur);  Kinston,  tributaries 
of  Fox  River. 

Vegetative  form:  Cysts,  round  or  elliptic,  not  over  1mm.  in  diameter, 
covered  by  resistant  membrane.    Color  whitish.    Polysporous. 

Spore:  Spatular,  approaching  roundish-oblong.  Shell  thin  and  trans- 
parent. Sutural  ridge  wide.  Two  polar  capsules,  pyriform,  of  equal  size. 
Sporoplasm  extending  forward  along  the  upper  surface.  Vacuole  could  not 
be  detected.  Dimensions:  length  14  to  17/i,  breadth  8.5/x,  thickness  5 
to  6/i. 

MYXOBOLUS  TRANSOVALIS  Gurley 
[Figs.  417  and  418] 

1893  Myxobolus  transovalis  Gurley  1893  :  415 

1894  Myxobolus  transovalis  Gurley  1894  :  242 

Habitat:  Under  scales  on  external  surface  of  Phoxinus  (Clinostomus) 
funduloides  Girard;  4  Mile  Run,  Carlius,  Va.,  tributary  of  Potomac  River 
(June),  No  fish  of  the  same  species  caught  from  the  same  locality  on 
August  29  of  the  same  year  was  found  infected. 

Vegetative  form:  It  is  not  certain  whether  cysts  exist  or  not.  Spores 
in  mass,  appear  to  be  held  together  by  a  small  gelatinous  or  mucoid  mass 


140  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [378 

which  has  no  attachment  to  the  subjacent  connective  tissue.  It  forms 
a  thin  discoidal  mass  situated  in  the  center  of  the  concave  surface  of  the 
scale.  The  color  of  the  mass  slightly  more  yellowish  than  the  surrounding 
tissue,  when  coagulated.  It  is  exceedingly  diflScult  to  detect  its  presence 
in  Vhe  fresh  state. 

Spore:  Elliptical,  with  the  largest  diameter  passing  thru  two  polar 
capsules.  Shell  thin.  Sutural  edge  narrow.  Two  polar  capsules  of  equal 
size  convergent.  Polar  filament  is  extruded  under  the  action  of  glycerine 
and  sulphuric  acid.  The  vacuole  in  the  sporoplasm  is  difficult  to 
detect.  Sporoplasm  also  contains  two  nuclei,  rarely  one,  1  to  1.5/*  in 
diameter.    Dimensions:  length  6  to  7/i,  breadth  Sju.  ^ 

MYXOBOLUS  OBESUS  Gurley 


[Figs. 

419  and  420] 

1883 

Balbiani 

1883 

:203 

1893 

Myxobolus  obesus 

Gurley 

1893 

:415 

1894 

Myxobolus  ?  obesus 

Gurley 

1894 

.239 

1899 

Myxobolus  obesus 

Labb^ 

1899 

:100 

1906 

Myxobolus  obesus 

C^pMe 

1906 

:60HS1 

Habitat:  On  Alburnus  alburnus  L.;  branchiae  and  kidney  of  A.  lucidus 
Heck.  (A .  jnirandella  Bl.);  Lac  du  Bourget. 

Vegetative  form:  Balbiani  gave  no  observation. 

Cepede  observed  as  follows:  Cysts,  ovoidal,  more  or  less  elongated 
or  variable  in  form,  not  exceeding  800ju  in  length.  In  kidney,  numerous  cysts 
were  of  subspherical,  ovoidal  or  rarely  irregularly  elongated  form.  Sub- 
spherical  cysts  500  to  600/x  in  average  diameter.    Polysporous. 

Spore:  Cepede  describes  as  follows:  Subcircular  or  ovoidal  in  front 
view;  lenticular  in  side  view.  Sutural  edge  exhibits  variable  numbers  (4 
to  5)  of  fold-like  markings  on  the  shell.  Polar  capsules  pyriform  and  of 
equal  size.  Coiled  polar  filament  distinct.  A  small  triangular  intercapsular 
appendix.  Sporoplasm  with  a  subspherical  and  clearly  outlined  vacuole 
and  two  nuclei.  Dimensions  in  vivo:  length  11.5  to  12/li,  breadth  7.5  to  8/i, 
thickness  5^.  Those  of  fixed  and  stained  spores:  length  11.25  to  11.50/*, 
breadth  7.25  to  7.50/*,  length  of  polar  capsule  5/t. 

Remarks:  Cepede  mentions  that  Alburnus  alburnus  L.  mentioned  by 
Gurley  is  "without  doubt"  identical  with  A.  lucidus  Heckel. 

MYXOBOLUS  CYCLOIDES  Gurley 


[Fig.  421] 

1841 

Miiller 

1841 

:  481,  486 

1893 

Myxobolus  cycloides 

Gurley 

1893 

:41S 

1894 

Myxobolus  cycloides 

Guriey 

1894 

:239 

1906 

Myxobolus  cycloides 

C6pede 

1906 

:  61-63 

1910 

Myxobolux  cycloides 

Wegener 

1910 

:  79-80 

379)  STUDIES  ON  MYXOSPORIDIA—KUDO  141 

Habitat:  Opercle,  pseudobranchiae  and  kidney  of  Leuciscus  ruHlus; 
branchiae  of  Scardinius  erythrophthalmus,  Blicca  bjorkna  L.,  Gohio  gobio 
L.,  Abramis  vintba  L.,  A.  brama  L.,  Rhodeus  amarus  Bl.,  Alburnus  alburnus 
L.,  Lota  lota  L.;  France  (Isere),  Germany  (Pregel,  Frisches  and  Kurisches 
Haff,  Masurische  See,  January,  May). 

Vegetative  form:  Wegener  observed  cysts  as  follows.  A  type:  1  to 
2mm.  by  0.4  to  0.7mm.  Form  exactly  like  that  of  Myxobolus  oviformis. 
B  type:  small  and  round,  present  in  groups.  C  type:  small  0.5mm.  by 
0.2mm. 

Spore :  Gurley  gave  the  following  short  diagnosis  from  the  observations 
of  J.  Miiller:  subcircular-ovate  or  broadly  rounded  elliptic,  length  12/*. 

Cepede  distinguishes  three  different  types  of  spores  as  follows :  Lenticu- 
lar in  side  view;  subcircular  (13.5/x  by  13/i),  oval  (14.7/i  by  11.4ju)  and 
ovoidal  (16/i  by  11^)  in  front  view.  Two  polar  capsules  of  equal  size  (6^ 
by  4jLi),  closely  set  or  separated  (3n  apart)  from  each  other.  Coiled  polar 
filament  distinct.  A  small  triangular  intercapsular  appendix.  Sporoplasm 
refractive  and  finely  granular.  Sutural  edge  exhibits  folds  of  variable 
number  at  the  posterior  portion.  Dimensions  of  fixed  and  mounted 
spores:  length  10.5  to  12/x,  breadth  7.5  to  S/x. 

Wegener,  without  noticing  Cepede's  paper,  also  mentions  three  different 
types  chiefly  distinguished  by  the  spore  as  follows: 

A  type  (common  form),  in  the  branchiae  of  Lota  lota,  Abramis  brama, 
A.  vimba,  Blicca  bjorkna,  Leuciscus  rutilus,  Alburnus  alburnus  and  scardi- 
nius erythrophthalmus .    Cysts  mentioned  above. 

Spore.  Rounded  or  oval;  flattened.  A  tail,  15/i  long,  was  noticed 
twice.  A  triangular  intercapsular  appendix.  Sutural  edge  usually  having 
folds.  Polar  capsules  often  differ  in  form  and  size  in  different  cysts,  tho 
they  are  constant  in  one  and  the  same  cyst,  causing  the  variability  in  size 
of  sporoplasm.  Dimensions:  length  11  to  12.5/i,  breadth  8  to  9ju,  polar 
capsule  4.5  to  6^  by  3  to  3.7/x,  in  many  cysts7.5/x  by  4jli. 

B  type.    In  the  fifth  gillarch  of  Gobio  gobio  L.  Cysts  mentioned  above. 

Spore.  Elongated  oval.  A  triangular  intercapsular  appendix.  Indis- 
tinct folds  on  sutural  edge.  Dimensions:  length  12.5  to  13. 5m,  breadth  8 
to  lOju,  polar  capsule  5  to  6^  by  3  to  4ju. 

C  type.  In  the  branchiae  of  Rhodeus  amarus  Bl.  and  Alburnus  alburnus 
L.  (April  and  May).     Cysts  mentioned  above. 

Spore.  Rounded.  Distinct  intercapsular  appendix.  Folds  distinct 
on  sutural  edge.  Dimensions:  length  12  to  15/x,  breadth  9  to  lOju,  polar 
capsule  5  to  7/x  by  3  to  4/i. 

MYXOBOLUS  SPHAERALIS  Gurley 

1874  Clapar^de  1874  :  113-114 

1893  Myxobolus  sphaeralis  Gurley  1893  :  415 

1894  Myxobolus  sphaeralis  Gurley  1894  :  240 


142  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [380 

Habitat:  Mucosa  of  branchiae  of  Coregonus  lavaretus  L.   (C.  fera); 
Lake  Geneva. 

Vegetative  form:  Cysts,  0.25  to  0.33mm.  in  diameter.    Polysporous. 
Spore:  Spherical,  9/z  in  diameter. 

MYXOBOLUS  ANURUS  Cohn 
[Figs.  422  and  423] 


1895 

Myxobolus  anurus 

Cohn 

1895 

:  42-43 

1896 

Myxobolus  anurtts 

Cohn 

1896 

:266 

1899 

Henneguya  psorospermica 

anura 

Labb£ 

1899 

:102 

1910 

Myxobolus  anurus 

Wegener 

1910 

:76 

1911 

Henneguya  psorospermica 

anura 

Nemeczek 

1911 

:146 

Habitat:  Branchiae  of  Esox  lucius  L.;  Konigsberg  (March,  December), 
Frisches  Haff,  Pregel,  Masurische  See,  Lotzen,  (September,  October). 

Vegetative  form:  Cysts  small  rounded  and  of  white  color.  Cohn 
measures  length  0.6mm., breadth  0.34mm.  Wegener's  form:  length  0.3 
to  0.5mm.  and  breadth  0.2  to  0.3mm. 

Spore:  Cohn's  descriptions  are  as  follows:  More  or  less  oval.  Dimen- 
sions: length  12  to  15/i,  breadth  4  to  6.8/*,  polar  capsule  5.5  to  7/x  by  2.1  to 
2.5/i,  length  of  polar  filament  32  to  38m. 

Wegener's  form :  Elongated  and  narrow,  often  with  a  tail.  Dimensions : 
length  15/i  (maximum  up  to  18^),  breadth  6  to  7/*,  polar  capsule  8n  by  3n. 

Remarks:  Tho  Labbe  classified  this  as  a  subspecies  of  Henneguya  pso- 
rospermica Thelohan,  Wegener's  observation  gives  stronger  basis  for  placing 
this  form  in  the  genus  Myxobolus. 

MYXOBOLUS  sp.  Gurley 
[Fig.  424] 

1882  B&tschli  1882  :  590 

1894        Myxobolus  sp.  incert.  Gurley  1894  :  214 

1899        Myxobolus  sp.  Labb6  1899  :  100 

Habitat:  Nais  lacustris  L.  {N.  prohoscidea);  Locality? 
Vegetative  form:  Cysts,  8mm.  by  4.25mm.    Polysporous. 
Spore:  Oval  or  circular;  tailed  or  untailed.    These  spores  of  diflrerent 
form  occur,  often,  without  order  in  the  same  cyst. 

MYXOBOLUS  sp.  Gurley 
[Fig.  425] 
1894        Myxobolus  sp.  incert.  Gurley  1894  :  239 

Habitat:  Body  cavity  of  Carassius  carassius  L.;  Leipsic. 
Vegetative  form:  Not  observed. 


381]  STUDIES  ON  MYXOSPORIDIA—KUDO  143 

Spore:  Broadly  elliptic;  shell  bivalve;  valves  equally  convex.  Sutural 
ridge.  Two  equal  polar  capsules.  Sporoplasm  with  a  vacuole.  Dimen- 
sions: length  14/i,  breadth  lO/i,  thickness  Six. 

Remarks:  This  species  seems  to  be  very  similar  to  M.  carassii  Kloka- 
6ewa  (page  150). 

MYXOBOLUS  sp.  Gurley 
[Figs.  426  to  429] 

1841  MiiUer  1841  :  480 

1894        Myxobolus  sp.  Gurley  1894  :  240-241 

1899        Myxobolus  sp.  Labb6  1899  :  100 

Habitat:  Skin  of  opercle,  in  the  branchiae,  on  the  head  or  on  the  fin  of 
Lucioperca  lucioperca  L.;  Germany,  Don. 

Vegetative  form:  Cysts  1.09  to  2.18mm.  in  diameter.  Color  whitish. 
Polysporous. 

Spore:  Rounded.  Thickness  equal  to  haK  the  breadth.  Sutural  ridge. 
Two  polar  capsules,  of  equal  size,  converging. 


MYXOBOLUS  CYPRINI  Doflein 
[Figs.  430  to  432] 

1896  Hofer  1896  : 2, 38-39 

1898  Myxobolus  cyprini  Doflein  1898  :  288,  320,  325 

1904  Myxobolus  cyprini  Hofer  1904  :  66-67 

1909  Myxobolus  cyprini  Doflein  1909  :  780-783 

1916  Myxobolus  cyprini  Doflein  1916  :  1026-1027 

Habitat:  Suppurative  connective  tissue  and  epithelium  of  kidney, 
liver  and  spleen  of  Cyprinus  carpio  L.,  rarely  Tinea  vulgaris  Cuv.  and 
Abramis  brama  L.;  Germany,  Austria.  According  to  Hofer  the  parasites 
cause  so-called  "small  pox  of  carp"  among  carp  in  German  waters. 

Vegetative  form:  Small  ameboid.  Form  irregular.  The  youngest 
form  with  a  single  or  many  nuclei,  is  found  in  the  epithelium  of  the  kidney. 
Multiplication  by  multiple  division,  the  nuclei  undergoing  amitotic 
division.  Endoplasm  contains  homogeneous,  yellow  and  refractive  bodies. 
Also  found  in  the  state  of  diffuse  infiltration.  Spores  are  found  in  the 
parenchym  of  the  kidney. 

Spore:  Oval.  Shell  thickened  (1.5)li  wide)  along  the  sutural  edge. 
Two  converging  polar  capsules  cross  each  other,  in  front  view,  at  the 
anterior  tip.  Sporoplasm  with  an  iodinophilous  vacuole.  Dimensions: 
length  21m,  breadth  ISjj.,  length  of  polar  capsule  6^i.  Doflein  (1916:1027) 
gives  the  following  dimensions:  length  10  to  16/x,  breadth  8  to  9/i. 

Hofer  gives  the  following  dimensions:  length  10  to  \2n  (up  to  16/i), 
breadth  8  to  \\n,  polar  capsule  5  to  6m  by  Zn,  sutural  edge  LSju. 


144  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [382 

MYXOBOLUS  NEUROBIUS  Schuberg  et  Schroder 
[Figs.  433  to  436] 
1905        Myxobolus  neurobius         Schuberg  and  Schroder  1905  :  49-56 

Habitat:  Nervous  tissue  of  TruUa  fario  L.;  Gutach  (May?). 

Vegetative  form:  Cysts,  usually  elongated,  often  spherical.  Elongated 
form  0.9mm.  by  0.02mm.  The  seat  of  the  cysts  is  between  the  medullary 
sheath  and  sheath  of  Schwann.  Neither  medullary  sheath  nor  axis-cylinder 
was  infected.  Cyst-membrane  could  not  be  made  out.  Cysts  contained 
only  full-grown  spores  without  any  younger  stage.    Polysporous. 

Spore:  Broad  oval  in  front  view;  spindle  shaped  in  side  view.  Anterior 
end  attenuated,  posterior  end  rounded.  Shell  somewhat  thick.  Sutural 
ridge  is  not  particularly  marked.  Edge  without  any  fold.  No  intercapsu- 
lar  appendix.  Sporoplasm,  with  a  large  and  spherical  iodinophilous 
vacuole  and  a  single  nucleus,  occupies  less  than  one  half  of  the  inner  space 
of  the  spore.  Two  polar  capsules,  pyriform,  fuse  into  one  at  the  anterior 
end.  Coiled  (8  to  10  times)  polar  filament  distinct.  Dimensions:  length 
10  to  12/i,  breadth  8ju,  thickness  6^,  polar  capsule  6  to  Tju  by  2jLt. 

MYXOBOLUS  AEGLEFINI  Auerbach 
[Figs.  437  to  441] 
1906        Myxobolus  aeglefini  Auerbach  1906  :  568-570 

1906  Myxobolus  aeglefini  Auerbach  1906a  :  115-119 

1907  Myxobolus  esmarkii  Johnstone  and     1907  :  204-208 

Woodcock 

1909  Myxobolus  aeglefini  Auerbach  1909  :  76-78 

1910  Myxobolus  aeglefini  Auerbach  1910c  :  181-182 

1911  Myxobolus  aeglefini  Nemeczek  1911  :  162 

Habitat:  Cartilage  and  bone  of  cranium  and  eye  of  Gadus  aeglefinis 
G.  callarias,  G.  merlangus  L.,  G.  morrhua  L.,  G.  esmarkii  and  Molva  vulgaris 
Flem.;  Norway,  Morecambe  (March). 

Vegetative  form:  Cysts  in  cartilage  and  bone  of  cranium  and  in  carti- 
lagineous  layer  of  the  sclerotic  of  the  eye.  Protoplasm  is  distinctly 
differentiated.  Ectoplasm  somewhat  vacuolated;  endoplasm  granular 
with  numerous  small  nuclei.    Polysporous. 

Johnstone's  observations  are  as  follows:  Round  the  peripheral  part 
of  the  cornea,  and  covered  loosely  by  conjunctiva  are  a  number  of  milk- 
white  rounded  or  oval  bodies,  from  about  1  to  3mm.  in  diameter.  Several 
of  these  fused  to  form  elongated  mass  which  lie  along  the  curvature  of  the 
periphery  of  the  eye.  These  cysts  also  invade  the  lateral  and  posterior 
parts  of  the  bulbus  ocuU.  In  sections,  the  cysts  lie  within  the  thickness 
of  cartilaginous  layer  of  the  sclerotic.  This  latter  is  enlarged  into  thick 
layer  (2mm.)  by  the  presence  of  the  cysts. 

Nemeczek  mentions  irregular  cysts  of  1.5mm.  in  diameter. 


383]  STUDIES  ON  MYXOSPORJDJA—KUDO  145 

Spore:  Elliptical  in  front  view.  Two  polar  capsules  convergent.  No 
intercapsular  appendix.  Sutural  edge  rather  thick  with  a  number  of  folds 
on  the  posterior  margin.  Sporoplasm  with  two  nuclei  and  an  iodinophilous 
vacuole.  Dimensions:  length  10.8  to  11.7/i,  breadth  9.9  to  10.4ju,  thick- 
ness 7.2  to  9n,  length  of  polar  capsule  4.5  to  5/x. 

Woodcock's  form  has  a  spore  with  the  following  characters: 
Slightly  ovoid.    Sporoplasm  always  contains  a  large  and  well  defined 
vacuole  and  two  nuclei.     Dimensions:  length  lO/x,  breadth  8m,  length  of 
polar  capsule  3.25  to  3.5ju. 

MYXOBOLUS  GIGAS  Auerbach 
[Figs.  442  to  445] 

1906        Myxoholus  gigas  Auerbach  1906  :  386-391 

1910        Myxobolus  gigas  Auerbach  1910c  :  182 

1912        Myxoholus  gigas  Parisi  1912  :  293-294 

Habitat:  Subcutaneous  connective  tissue  of  the  operculum  of  Abramis 
hrama  L.;  Karlsruhe,  Pavia.  Parisi  observed  cysts  on  the  side,  on  the 
caudal  fin  (5  cysts  on  rays),  on  other  fins,  branchiae  and  in  the  internal 
organs  of  the  fish. 

Vegetative  form:  Cysts,  spherical  or  ovoidal.  No  cyst  membrane 
composed  of  the  connective  tissue  of  the  host.  Protoplasm  is  indistinctly 
differentiated.  Ectoplasm  thin  and  radially  striated,  which  gradually 
turns  into  endoplasm.  Endoplasm  finely  granular,  contains  numerous 
nuclei  (2.5  to  2.7/i  in  diameter).  Size  of  greatest  form  360ju  by  290  to  300/i. 
According  to  Parisi  size  up  to  1.5mm. 

Spore :  Elliptical  when  viewed  from  the  front.  Sutural  edge  somewhat 
narrow,  having  a  number  of  folds  at  the  posterior  portion.  Sporoplasm 
with  an  iodinophilous  vacuole  and  two  nuclei.  Dimensions:  length  16.9  to 
21.6m,  breadth  13  to  16. 2^  thickness  9ju,  length  of  polar  capsules  7.8/i, 
length  of  polar  filament  90m  (sulphuric  acid) . 

Parisi  gives  150m  for  the  length  of  polar  filament. 

MYXOBOLUS  VOLGENSIS  Reuss 

[Figs.  446  to  448] 

1906        Myxoholus  volgensis  Reuss  1906  :  200-201 

Habitat:  Branchiae,  cornea  and  dorsal  fin  of  Lucioperca  volgensis 
Pall;  Volga. 

Vegetative  form:  Cysts,  spherical,  0.3  to  1mm.  in  diameter.  Poly- 
sporous. 

Spore:  Broad  elliptic  or  rounded.  Sutural  edge  has  at  least  3  folds. 
Sporoplasm  with  an  iodinophilous  vacuole.  Dimensions:  length  8.25  to 
9.5m,  breadth  7.25  to  8.25m,  thickness  4.5  to  5.5m,  polar  capsule  Sp,  by  Ipi. 


146  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [384 

MYXOBOLUS  SCARDINII  Reuss 
[Fig.  449] 
1906        Myxobolus  scardinii  Reuss  1906  :  201 

Habitat:  Branchiae  of  Scardinius  erythrophthalmus  L. ;  Volga. 
Vegetative  form:  Cysts,  elongated  oval.    Smaller  cysts  rounded  oval, 
0.8mm.  by  0.5mm.,  the  larger  forms  elongated,  1.2mm.  by  0.5mm.    Poly- 
sporous. 

Spore:  Broad  elliptical.  Sutural  edge  narrow,  having  folds.  A  larger 
triangular  intercapsular  process.  An  iodinophilous  vacuole  in  sporoplasm. 
Dimensions:  length  11  to  i2fi,  breadth  9  to  9.5ju,  thickness  4.5  to  5^, 
polar  capsules  5n  by  2.5/i. 

MYXOBOLUS  PHYSOPHILUS  Reuss 
[Figs.  450  and  451] 
1906        Myxobolus  physophilus  Reuss  1906  :  201-202 

Habitat:  Surface  of  air  bladder  of  Scardinius  erythrophthalmus  L.; 
Volga. 

Vegetative  form:  Cysts,  rounded,  1.5mm.  in  diameter.    Polysporous. 

Spore:  Oval,   with  attenuated  anterior  end.     Sutural  edge  narrow 

and  smooth.    Polar  capsules  rather  large.    An  iodinophilous  vacuole  in 

sporoplasm.    Dimensions:  length  12  to  13/i,  breadth  8.25  to  9/i,  thickness 

6.5  to  7/i,  polar  capsules  6/i  by  2.5/*. 

MYXOBOLUS  MACROCAPSULARIS  Reuss 
[Fig.  452] 
1906        Myxobolus  macrocapsularis       Reuss  1906  :  202 

Habitat:  Branchiae  of  Blicca  bjorkna  L.;  Volga. 

Vegetative  form:  Cysts,  Elongated  oval.  Size:  1mm.  by  0.5mm. 
Polysporous. 

Spore:  Oval  with  greatly  attenuated  anterior  portion.  Sutural  edge 
broad  and  without  any  fold.  Polar  capsules  rather  large.  An  iodinophil- 
ous vacuole  in  sporoplasm.  Dimensions:  length  11  to  13;*,  breadth  8.25  to 
9.25/*,  thickness  5.5/*,  polar  capsules  6/*  by  2.5  to  3/i. 

MYXOBOLUS  SANDRAE  Reuss 

[Fig.  453] 

1906        Myxobolus  sandrae  Reuss  1906  :  202-203 

Habitat:  Muscle  of  Luciop&rca  sandra  Cuv.;  Volga. 

Vegetative  form:  Cysts.  Rounded,  0.5mm.  in  diameter.  Polyspor- 
ous. 

Spore:  Oval.  Sutural  edge  broad  with  many  distinct  folds.  An 
iodinophilous  vacuole  in  sporoplasm.  Dimensions:  length  9.25  to  10/*, 
breadth  7.25  to  8.25/*,  thickness  4  to  5/*,  polar  capsules  3.5/i  by  2/*. 


3851  STUDIES  ON  MYXOSPORIDIA— KUDO  147 

MYXOBOLUS  BRAMAE  Reuss 

[Fig.  454] 

1906        Myxobolus  bramae  Reuss  1906  :  203-204 

Habitat:  Branchiae  of  Abramis  brama  L.;  Volga. 

Vegetative  form:  Cysts.  Oval,  O.Smm.  long,  0.25mm.  broad.  Poly- 
sporous. 

Spore:  Oval  to  nearly  spherical.  Sutural  edge  narrow  and  with  indis- 
tinct folds.  Two  polar  capsules,  with  a  small  triangular  intercapsular 
process.  An  iodinophilous  vacuole.  Dimensions:  length  11  to  12/*, 
breadth  9.25  to  10/x,  thickness  4.5  to  5.5/i,  polar  capsules  4  to  5/i  by  2.25/*. 

MYXOBOLUS  CYPRINICOLA  Reuss 
.     [Fig.  456] 
1906        Myxobolus  cyprinicola  Reuss  1906  :  204 

Habitat:  Branchiae  of  Cyprinus  carpio  L.;  Volga. 

Vegetative  form:  Cysts,  oval,  0.5mm.  by  0.3mm.    Polysporous. 

Spore:  Elongated  oval.  Sutural  edge  narrow  with  many  indistinct 
folds.  An  iodinophilous  vacuole.  Dimensions:  length  9.25  to  10/t,  breadth 
7  to  7.25/x,  thickness  5  to  5.5/*,  polar  capsules  4.5/*  by  2.5  to  3/*. 

MYXOBOLUS  BALLERI  Reuss 

[Fig.  455] 
1906        Myxobolus  balleri  Reuss  1906  :  204-205 

Habitat:  Branchiae  of  Abramis  ballerus  L.;  Volga. 

Vegetative  form:  Cysts.    Elongated,  1.5mm.  by  0.5mm.    Polysporous. 

Spore:  Oval,  slightly  pointed  at  the  anterior  end.  A  triangular 
intercapsular  appendix.  Sutural  edge  smooth.  An  iodinophilous  vacuole. 
Dimensions:  length  11  to  12/*,  breadth  9.25  to  10/*,  thickness  5.5  to  6.5/*, 
polar  capsules  5.5/t  by  2.75/*. 

MYXOBOLUS  SQUAMAE  Keysselitz 
[Figs.  457  to  459] 
1908        Myxobolus  squamae  Keysselitz  1908  :  273-274 

Habitat:  Inner  surface  of  the  scales  oi  Barbus fluviatilis  Agass.;  Mosel 
and  Neckar. 

Vegetative  form:  Form  variable;  rounded,  oval,  elongated  or  rarely 
branched.  The  outline  of  the  body  is  not  smooth  but  irregular  with  numer- 
ous small  tooth-like  projections  with  which  the  body  comes  in  contact 
with  the  surrounding  substance.  The  parasites  seem  to  be  able  to  dissolve 
the  substance  composing  the  scale.  Length  50  to  800/*.  In  one  scale,  one 
or  many,  up  to  8,  individuals  were  found.  All  showed  only  advanced  stages 
of  spore  formation.  The  parasites  are  surrounded  by  a  variously  developed 
envelope  of  connective  tissue.    Polysporous. 


148  ILLINOIS  BIOLOGICAL  MONOGRAPHS  (386 

Spore:  Elongated  oval.  Two  polar  capsules,  with  7  to  8  times  coiled 
polar  filament.  A  triangular  intercapsular  projection.  Sporoplasm  with 
an  iodinophilous  vacuole.  Dimensions:  length  10  to  10.5)li,  breadth  8  to 
8.5m,  length  of  polar  capsule  4.5^. 

MYXOBOLUS  CORDIS  KeysseUtz 

[Figs.  460  and  461] 

1908        Myxobolus  cordis  Keysselitz  1908  :  279-282 

Habitat:  Muscle  of  ventricle,  rarely  that  of  bulbus  arteriosus  of  Barbus 
fluviatilis  Ag.,  spores  found  in  kidney,  liver  and  spleen  in  the  condition  of 
somewhat  scattered  infiltration;  Germany  (Mosel  and  Neckar). 

Vegetative  form:  Elongated,  oval,  sausage  or  club  form.  The  body 
whitish,  later  yellowish.  Size  from  0.25  up  to  4mm.,  usually  1  to  1.5mm.  in 
length.  Propagative  stage  and  cysts  observed.  One  end  of  the  body  is 
held  more  or  less  deeply  in  the  muscle  and  is  covered  by  cellular  envelope 
as  in  Myxobolus  tnusculi,  while  remaining  larger  portion  of  the  body  is  sus- 
pended freely  inside  of  the  ventricle,  covered  with  a  thin  layer  probably  of 
endocardiac  cells.  Fish  30  to  45cm.  long  harboured  40  to  60  parasites. 
No  movements.  Ultimately  the  cysts  are  formed  with  differentiated 
protoplasm.     Polysporous. 

Spore:  Oval.  Shell  very  thin  at  the  anterior  end.  At  the  posterior 
end,  cell-like  appendage,  2  to  3n  wide  which  is  probably  formed  by  both 
valves,  is  present.  Two  pyriform  polar  capsules  at  the  anterior  end,  which 
show  the  polar  filament  coiled  7  to  8  times.  Sporoplasm  with  a 
comparatively  large  and  oval  iodinophilous  vacuole  and  two  nuclei, 
rarely  one  (syncaryon).  Dimensions:  length  12/i,  breadth  10/x,  length  of 
polar  capsule  4.5)u. 

MYXOBOLUS  MUSCULI  Keysselitz 

[Figs.  462  to  464] 

1908        Myxobolus  tnusculi  Keysselitz  1908  :  282-286 

Habitat:  Muscle  of  the  main  body,  rarely  that  of  fins  and  operculum, 
B.nd  kidney  oi  Barbus  Jluviatilis  AgSLSS.  of  various  size  (youngest  fish  found 
infected,  2  months  old),  spores  in  liver,  spleen,  kidney  and  ovary  (not  the 
ovum)  in  diffuse  infiltration;  Mosel  and  Neckar. 

Vegetative  form:  Elongated.  Body  whitish  opaque,  with  differentiated 
protoplasm.  Smallest  individual  observed,  24/i.  Large  form  2mm.  in 
length.  Many  trophozoites  are  found  closely  situated,  forming  a  large 
mass  of  parasites  that  reached  dimensions  of  4mm.  by  2mm.  The 
surrounding  envelope,  varying  in  thickness,  composed  of  cells  with 
elongated  nuclei  as  those  of  perimysium.  Young  cysts  surrounded  by  thin 
layer  of  ectoplasm.    Polysporous. 


3871  STUDIES  ON  MYXOSPORJDIA—KUDO  149 

Spore:  Oval.  Two  polar  capsules  usually  unequal.  Shell  as  in  M. 
cordis  with  a  small  peg  closer  to  the  anterior  end,  polar  filament  coiled  4  to 
5  times,  visible  in  the  capsule.  Sporoplasm  with  rarely  one  (syncaryon), 
but  usually  two  nuclei  and  an  iodinophilous  vacuole.  A  posterior  process 
as  is  seen  in  the  spores  of  M.  cordis,  but  much  smaller,  was  occasionally 
observed.  Dimensions:  length  11/i,  breadth  Sn,  polar  capsules  6jLt  and 
4/i  long. 

MYXOBOLUS  sp.   Miyairi 

1909  Myxobolus  sp.  Miyairi  1909  :  126 
Habitat:  Branchiae   of   loach    {Misgurnus   anguillicaudatus    Cant.?); 

Fukuoka?  (Nippon). 

Vegetative  form:  Cysts  were  not  observed. 
Spore:  No  description. 

MYXOBOLUS  sp.  Wegener 
[Fig.  465] 

1910  Myxobolus  sp.  Wegener  1910  :  78 
Habitat:  Branchiae  (gill-arch)  of  Percafluviatilis  L.;  Germany  (Frisches 

Haff,  March).     Only  one  case. 

Vegetative  form:  Cysts  on  a  gill-arch,  white  and  round,  with  a  diameter 
of  1.1mm.     Polysporous. 

Spore:  Form  and  size  very  variable.  Rounded  or  elliptical,  pointed  at 
the  anterior  end.  Sutural  edge  showing  folds  at  the  posterior  portion. 
Dimensions:  length  8  to  10)u  (in  round  form)  and  Wjx  (in  elliptical  form), 
breadth  8  to  9/i,  polar  capsules  4  to  5ju  by  2  to  3/x,  length  of  polar  filament 
40m. 

MYXOBOLUS  PERMAGNUS  Wegener 
[Fig.  466] 

1910  Myxobolus  permagnus  Wegener  1910  :  78-79 
Habitat:   Branchiae  and  operculum  of  P^rca /wwa^^Vu  L.,  air  bladder 

of  Scardinius  erythrophthalmus  L.;  Konigsberg  (May),  Pregel  (March). 

Vegetative  form:  Cysts  rounded  in  form  and  white  in  color,  resemble 
to  those  of  M.  gigas.  No  clear  ectoplasm  layer,  nor  typical  protoplasmic 
structure.     Polysporous. 

Spore:  Oval,  sharply  pointed  at  the  anterior  end.  Sutural  edge  with 
5  to  6  distinct  folds  at  the  posterior  portion.  Polar  filament  visible  in' the 
polar  capsules.  Dimensions:  length  17  to  18/x,  breadth  10  to  13/:,  polar 
capsules  7  to  8/x  by  3.5  to  4ju. 

MYXOBOLUS  ROTUNDUS  Nemeczek 
[Fig.  467] 

1911  Myxobolus  rotundus  Nemeczek  1911:156-157 
Habitat:  Branchiae  of  Abramis  brama  L.;  Austria. 


ISO  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [388 

Vegetative  form:  Cysts,  ovoidal  or  spindle  form,  1  to  3mm.  long  and 
1  to  1.5mm.  wide.  Body  white.  An  extraordinary  large  number  of  spores 
were  found  in  the  cysts.    Polysporous. 

Spore:  Round  or  slightly  oval,  when  viewed  from  the  front.  Greatly 
flattened  in  side  view.  Polar  capsules  convergent,  with  no  intercapsular 
body.  Shell  smooth.  Sutural  edge  narrow,  without  folds.  Dimensions: 
length  lO/i,  breadth  9,8/i,  thickness  3/x,  polar  capsules  3.8  to  5/x  long,  length 
of  polar  filament  40jtt. 

MYXOBOLUS  MINUTUS  Nemeczek 
[Fig.  468] 

1911  Myxobolus  mintUus  Nemeczek  1911  :  160 
Habitat:  Branchiae  of  Leuciscus  sp.;  Austria. 

Vegetative  form:  Cysts  spherical,  oval  or  elongated  with  white  color. 
Size:  0.5  to  3mm.  by  0.5  to  1mm.    Polysporous. 

Spore:  Rounded  oval,  similar  to  that  of  Myxobolus  rotundus.  Shell 
smooth.  Sutural  edge  narrow  without  folds.  Sporoplasm  with  an  iodino- 
philous  vacuole.  No  intercapsular  appendix.  Dimensions:  length  6m, 
breadth  4.2  to  5ju,  polar  capsule  ^n  by  2/i,  length  of  polar  filament  50  to  60, 
often  70/Lt. 

MYXOBOLUS  sp.  Lebzelter 

1912  Myxobolus  sp.  Lebzelter  1912  :  296-297 
Habitat:  Gall-bladder  of  Thymallus  thymallus  L. 
Vegetative  form:  Not  observed. 

Spore:  Sutural  ridge  distinct.    Dimensions,  length  5^,  breadth  3^. 

MYXOBOLUS  MAGNUS  Awerinzew 
[Figs.  469  and  470] 

1913  Myxobolus  magnus  Awerinzew  1913  :  75-76 
Habitat:  Eye  of  Acerina  cernua  L.;  Petrograd. 

Vegetative  form:  Trophozoites  form  white  spots  in  the  tissue  of  iris, 
with  many  spores  (300  to  400).  Each  pansporoblast  forms  in  most  cases 
two,  sometimes  3  or  5  spores!    Polysporous. 

Spores:  Large,  elongated  roundish,  slightly  flattened.  Sutural  edge 
somewhat  thick,  forming  a  wide  ridge,  with  4  to  5  folds  at  the  posterior 
portion.  Polar  capsules  do  not  cross  each  other.  Sporoplasm  with  an 
iodinophilous  vacuole  and  two  nuclei.  Dimensions:  length  38  to  45m, 
breadth  32  to  38m,  thickness  28  to  35m,  length  of  polar  capsules  15  to  17m, 
diameter  of  the  vacuole  12  to  16m. 

MYXOBOLUS  CARASSII  Klokacewa 
[Figs.  471  to  473] 

1914  Myxobolus  carassii  Klokacewa  1914  :  182-184 
Habitat:  Body  cavity,  liver  and  intestine  of  Carassius  vulgaris  L.; 

Petrograd? 


389]  STUDIES  ON  MYXOSPORJDJA—KUDO  151 

Vegetative  form:  Cysts  spherical.  Those  in  liver  and  intestine  yellowish, 
surrounded  by  an  envelope  composed  of  fibrous  connective  tissue.  Second- 
ary cysts  are  formed.    Polysporous. 

Spore:  Oval,  in  front  view.  Two  ovoidal  polar  capsules  convergent 
at  the  slightly  attenuated  anterior  end.  Coiled  polar  filament  visible. 
Sporoplasm  with  an  iodinophilous  vacuole  and  two  nuclei.  Sutural  edge 
shows  folds  in  some  cases.  Dimensions:  length  13  to  17ju,  breadth  8  to 
10/i,  thickness  5  to  In,  polar  capsules  6  to  Ifx  long. 

Remarks:  Compare  with  Myxobolus  sp.  Gurley  on  page  142. 

MYXOBOLUS  sp.  Southwell 
1915        Myxobolus  sp.  Southwell  1915  :  312-313 

Habitat:  Subcutaneous  intermuscular  tissue  of  Rasbora  (Cyprinus) 
daniconius  Day;  from  a  stream  near  Katwan,  Mirzapore  (U.P.),  India. 

Vegetative  form:  6  cysts  found  on  four  fish.  The  seat  is  immediately 
below  the  scales,  in  the  epidermis.  Color  milky  white.  Soft,  flattened  and 
roughly  oval  in  shape.  Greatest  length  found,  1.1mm,  No  pigment  was 
present  on  the  cyst. 

Spore :  Two  equal  capsules,  with  a  very  short  tail-like  process.  Sporo- 
plasm with  vacuole;  iodine  treatment  could  not  be  carried  out. 
Dimensions:  length  13^,  breadth  I3fi,  polar  capsule  4)u  by  4/i(?). 

Remarks:  Dimensions,  especially  that  of  polar  capsule  seem  to 
be  misprinted.  Southwell  gave  one  figure  of  a  fish  with  a  cyst  near  the 
dorsal  fin.  He  thinks  that  "it  is  quite  possible  that  our  parasites  may 
belong  to  Myxobolus  cyprini."  The  incomplete  observation  without  any 
figure,  leads  the  writer  to  leave  the  form  also  as  Myxobolus  sp.  Southwell. 

MYXOBOLUS  FUNDULI  Kudo 

[Figs.  474  to  476] 

1915        Myxobolus  musculi  Hahn  1915  :  201-205 

1917        Myxobolus  musculi  Hahn  1917  :  91-104 

Habitat:  Branchiae  and  muscle  of  Fundulus  heteroclitus,  F.  majalis; 
Woods  Hole.  Hahn  claims  that  he  succeeded  in  causing  experimental 
infection  in  F.  diaphanus  and  Cyprinodon  variegatus  by  inoculation. 

Vegetative  form:  Hahn  uses  quite  a  number  of  different  terms  from 
those  that  are  ordinarily  used  in  describing  Myxosporidia,  without  giving 
any  definitions.  Naturally  it  is  hard  to  put  what  he  wrote  in  several  pages 
in  the  following  lines.  Granular  vegetative  forms  produce  a  great  many 
pansporoblasts,  each  with  a  single  spore.  "Trophoplasm"  is  difficult  to 
stain.  Size:  74  by  33n,  24  by  I9fi.  Cysts  within  and  between  the  muscle 
fibers,  containing  several  hundred  spores. 

Spore:  Hahn's  descriptions  may  be  summarized  as  follows:  Dimen- 
sions: length  14.3^,  breadth  6.7/x,  thickness  6.7/t  to  2/3  of  width,  polar 


152  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [390 

capsule,  6.5/x  by  2n,  polar  filament  3  to  4  times  the  length  of  the  spore 
(42.9  to  57.2)Li).  Polar  filament  coiled  10  to  14  times.  Shell  thin,  almost 
invisible.  The  spores  found  in  the  gill:  length  12  to  13.4ju,  breadth  6/i  to 
10.4ai.    a  vacuole  is  present  in  the  sporoplasm. 

Remarks:  Examination  of  Hahn's  first  paper  suggested  that  he  was 
dealing  with  the  present  form  as  a  new  species  tho  he  did  not  mention 
at  all  Keysselitz  who  gave  the  name  Myxoholus  musculi  Keysselitz  to 
the  parasite  of  Barbus  fiuviatilis  from  German  riverg.  I  was  informed  by 
Hahn  that  he  gave  the  name,  Myxoholus  musculi,  without  knowing  the 
fact  that  it  was  preoccupied  by  Keysselitz  (1908)  (see  page  148)  and  that 
tho  he  became  aware  of  it  later,  he  can  not  determine  differences  by  which 
the  two  forms  can  be  distinguished.  A  comparison  of  the  descriptions 
of  Keysselitz  and  Hahn,  however,  shows  that  these  two  forms  differ  in 
several  respects.  Hence  the  latter  form  is  recorded  here  as  a  distinct 
species  under  the  new  name. 

It  is  interesting  to  note  that  very  similar  forms,  one  without  an  iodino- 
philous  vacuole  at  any  stage  of  spore-formation  (Myxosoma  funduli  Kudo, 
see  page  125)  and  the  other  with  a  vacuole,  occur  in  the  same  hosts 
in  the  same  locality.  As  mentioned  above,  the  reader  is  requested  to  refer 
to  Hahn's  original  paper  for  further  data. 

.  MYXOBOLUS  PLEURONECTIDAE  Hahn 

[Fig.  477] 

1917        Myxoholus  pleuronectidae  Hahn  1917  :  160-161 

Habitat:  Subcutaneous  muscular  tissue  of  Pseudopleuronectes  ameri- 
canus;  Woods  Hole. 

Vegetative  form :  Similar  to  that  of  Myxoholus  f unduli. 

Spore:  Hahn  writes  as  follows:  Shape  and  appearance  resembles 
Myxoholus  pfeifferi.  Dimensions:  length  14.5/x,  breadth  11.9ju,  polar 
capsules  6n  by  3.7 fx. 

MYXOBOLUS  CAPSULATUS  Davis 

[Fig.  478] 
1917        Myxoholus  capsulatus  Davis  1917  :  237 

Habitat:  Visceral  connective  tissues  of  Cyprinodon  variegatus;  Beaufort. 

Vegetative  form:  Irregular  form.  In  the  state  of  diffuse  infiltration. 
Polysporous. 

Spore:  Pyriform,  flattened.  Polar  capsules  large  and  pyriform,  filling 
almost  entire  cavity  of  the  spore.  Sporoplasm  relatively  small.  lodino- 
philous  vacuole  visible  in  living  spore.  Dimensions:  length  16)li,  breadth 
10  to  1  Iju,  polar  capsules  1 1/x  by  4/i,  length  of  polar  filament  84/i. 


3911  STUDIES  ON  MYXOSPORJDJA—KUDO  153 

MYXOBOLUS  NODULARIS  Southwell  et  Prashad 

[Figs.  479  and  480] 

1918        Myxobolus  nodularis        Southwell  and  Prashad        1918  :  347 

Habitat:  In  the  muscles  of  Rashora  daniconius  occurring  in  two  fish 
on  the  sides,  and  in  another  as  a  globular  cyst  near  the  anus;  Mirpur, 
Dacca  (June).    Type  specimens,  numbered  P  52/1, 

Vegetative  form:  Cysts  rounded  or  slightly  elongated,  varying  in  length 
3.5  to  3.8mm.  and  2.3  to  2.8mm.  in  width.  Creamy  yellow  in  color,  in  one 
case  appearing  blackish  owing  to  the  large  number  of  black  granules 
scattered  in  its  surface. 

Spore:  Ovoidal.  Sutural  ridge  very  wide  (about  1/5  thickness  of  the 
spore).  Two  polar  capsules  of  equal  size,  which  show  coiled  polar  filaments 
clearly.  Dimensions:  length  9/i,  breadth  7.2/i,  length  of  polar  capsule 
3.4;ti,  that  of  polar  filament  18.3/i. 

MYXOBOLUS  HYLAE  Johnston  et  Bancroft 
[Figs.  591  to  593] 


1888 

Fletcher 

1888 

:337 

1890 

Haswell 

1890 

:661 

1909 

Myxobolus  sp. 

Johnston 

1909 

:29 

1910 

Myxobolus  sp. 

Cleland  and 

Johnston 

1910 

:25 

1918 

Myxobolus  hylae 

Johnston  and 

Bancroft 

1918  ; 

:  171-175 

Habitat:  In  the  testes,  vasa  efferentia  and  oviducts  of  Hyla  aurea; 
Sidney,  Australia  (April,  other  months  not  mentioned).  Fletcher  observed 
the  parasites  also  in  the  urinary  bladder  of  both  sexes,  which  fact  was  not 
confirmed  by  Johnston  and  Bancroft  on  account  of  the  scarcity  of  the 
material.  The  latter  authors  could  not  infect  Hyla  caerulea  by  feeding 
infected  testes  or  the  cysts,  giving  the  conclusion  that  the  parasite  is 
specific  to  H.  aurea.  The  male  is  more  often  attacked  by  the  parasite 
than  the  female.  The  infected  animal  appeared  sickly  and  emaciated. 
As  to  the  infection  in  kidneys,  they  write  as  follows:  In  one  male  specimen 
both  testes  and  both  kidneys  were  affected,  and  the  upper  parts  of  the 
ureters  adjacent  to  the  kidneys  were  swollen  and  milky  in  appearance. 
In  another,  in  addition  to  the  testes,  the  adjacent  kidney  and  mesentery 
were  attacked.  No  spores  have  yet  been  detected  by  them  in  sections  of 
the  kidney  tubules. 

Vegetative  form:  Johnston  and  Bancroft  describe  as  follows: 
Cysts:  in  male,  either  imbedded  in  the  tissue  or  may  project  freely  into 
the  coelom  of  the  testes;  in  female,  lying  between  the  layers  of  the  wall, 
being  projected  into  the  lumen  of  the  oviduct.     Size  from  those  of  micro- 
scopic dimensions  up  to  2  to  3mm.  in  diameter.  In  sections,  the  protoplasm 


154  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [392 

is  differentiated  into  two  regions.  The  outer  layer  (ectoplasm)  surrounds 
the  body  as  a  thin,  light-staining  region,  while  the  endoplasm  being  den- 
ser and  of  more  or  less  granular  structure  filled  with  spores  especially  in 
the  central  portion. 

Spore:  Johnston  and  Bancroft  describe  as  follows: 

Form  somewhat  variable,  caused  by  the  reduction  in  length.  Oval, 
egg-shaped  or  nearly  circular  in  front  view.  Sutural  ridge  slightly  thick- 
ened. Two  pyriform  polar  capsules  are  located  at  the  anterior  end. 
Sporoplasm  with  an  iodinophilous  vacuole  (2ju  in  diameter),  shows  usually 
two  distinct  nuclei,  rarely  but  one.  Dimensions:  length  variable,  diameter 
of  circular  form  7  to  8/x,  breadth  8  to  lOju,  thickness  about  6/x,  thickness  of 
shell  1/x,  polar  capsules  4  to  5/i  by  2/i,  length  of  polar  filament  90  to  98ju 
(acids  or  alkalies).  , 

MYXOBOLUS  AUREATUS  Ward 
[Figs.  643  to  649] 
1919        Myxoholus  aureatus  Ward  1919  :  49 

Habitat:  Between  the  ectodermal  layers  of  the  fin  membrane  of 
Notropis  anogenus',  Put-in-Bay,  Lake  Erie  (August).  Out  of  thirty  fish, 
two  to  three  cm.  in  length,  seven  were  found  to  be  infected.  The  infected 
fish  were  not  inferior  in  size  or  vigor  to  others  of  the  same  species.  The 
most  heavily  infected  one  was  the  most  vigorous  of  all.  The  number  of 
cysts,  in  the  individual  fish,  varied  from  one  to  forty,  being  confined  in  the 
fin.  The  cysts  are  always  separated  from  each  other,  tho  in  a  few  instances 
they  were  apparently  connected. 

Vegetative  form:  The  parasite  forms  cysts  between  the  ectodermal 
layers  of  the  fin  membrane.  The  cyst  is  a  smooth  margined  ellipsoid, 
measuring  from  1  to  1.6  mm.  in  layer  diameter  and  from  0.8  to  1.2  mm. 
along  its  transverse  axis.  The  opaque  cyst  is  of  a  clear  orange  yellowish 
color.  This  gilt  color  is  contained  in  the  cyst  wall,  fading  away  in  alcohol 
and  formol.  The  chromatophores  of  the  skin  of  the  host  are  distinctly 
more  abundant  on  the  cyst  than  in  other  parts  of  the  skin,  and  the  older 
the  cyst  the  more  abundant  the  chromatophores.  The  wall  of  the  cyst  is 
noticeably  tough  and  thick.  In  section,  the  protoplasm  shows  a  poor 
differentiation  into  ectoplasm  and  endoplasm.  The  former  granular  and 
reticular,  covers  the  entire  surface  as  a  thin  layer,  while  the  latter  is  highly 
vacuolated,  containing  only  mature  spores.     Polysporous. 

Spore:  Ovoid;  slightly  pointed  anterior  and  rounded  posterior  ends  in 
front  view;  slightly  compressed  in  lateral  view.  Sutural  ridge  distinct. 
The  shell  is  of  moderate  thickness,  and  bears  a  flange  at  the  posterior 
half  in  some  spores.  Two  pyriform  polar  capsules,  frequently  of  slightly 
different  dimensions,  are  at  the  anterior  part  of  the  spore.  No  intercapsu- 
lar  appendix  is  present.     When  the  spore  is  allowed  to  stand  for  24  hours 


393]  STUDIES  ON  MYXOSPORIDIA—KUDO  ISS 

or  more  in  water,  the  polar  filaments  are  extruded.  The  binucleated 
finely  granular  sporoplasm  shows  an  iodinophilous  vacuole.  Dimensions: 
length  12.4  to  13.5/i,  breadth  6.5  to  7.5/*,  thickness  5/i,  length  of  polar 
capsule  6  to  7/i  (rarely  7.5/x),  length  of  polar  filament  about  20  to  26/i, 
diameter  of  iodinophilous  vacuole  about  2n. 

MYXOBOLUS  MIYAIRII  nov.  spec. 
[Fig.  481] 
1909        Myxoholus  sp.  Miyairi  1909  :  130,  131-132 

Habitat:  Intestinal  wall  of  Parasilurus  asotus  L.;  Fukuoka  ?  (Nippon) 

Vegetative  form:  Cysts.  Size  rather  small  up  to  0.5mm.  Full-grown 
spores  as  well  as  those  in  developmental  stages  fill  the  central  portion  of 
cysts,  while  numerous  nuclei  are  chiefly  found  along  the  periphery  of  end- 
plasm. 

Spore:  Elongated  elliptic.  Two  polar  capsules  of  nearly  same  size. 
Sporoplasm  with  a  comparatively  large  iodinophilous  vacuole.  Dimen- 
sions: length  13  to  14.5)u,  breadth  6  to  7/x,  length  of  polar  capsules  4.5iLt, 
length  of  polar  filament  30  to  35/i. 

Remarks.  As  the  descriptions  show  the  form  and  structure  are  dis- 
tinguishable from  other  species,  the  writer  establishes  the  present  species. 

MYXOBOLUS  KOI  nov.  spec. 
[Figs.  482  to  485] 

Habitat:  In  the  connective  tissue  of  the  gill  filament  of  CypHnus  carpio 
L.;  Tokio  (April).    One  fish  was  found  infected  in  a  slight  degree. 

Vegetative  form:  Cysts  small  and  spherical;  white  in  color.  Size  up  to 
230m  in  largest  diameter.  The  seat  similar  to  Myxoholus  toyamai.  The 
structure  of  the  cysts,  observed  in  section  preparations,  is  also  similar  to 
the  above  mentioned  unicapsular  Myxoholus. 

Spore:  Oval  with  attenuated  anterior  and  rounded  posterior  ends  in 
front  view;  elongated  pyriform  in  side  view.  Shell  comparatively  thin.  No 
marking  on  shell.  Sutural  ridge  fairly  well  marked.  No  intercapsular 
appendix.  Two  polar  capsules  are  pyriform,  large,  and  of  usually  equal 
form  and  size.  Coiled  polar  filament  distinct  in  vivo.  Sporoplasm  rather 
small,  finely  granular,  shows  two  nuclei  in  almost  all  spores.  An  iodino- 
philous vacuole  is  deeply  stained  by  Lugol's  solution.  Dimensions:  14  to 
16m,  breadth  8  to  9fx,  thickness  5  to  6m,  polar  capsule  8  to  9m  by  2.5  to  3m, 
length  of  polar  filament  72m  in  average  (KOH). 

MYXOBOLUS  ORBICULATUS  nov.  spec. 
[Figs.  566  to  576] 

Habitat:  Muscle  of  myotomes  of  Notropis  gilherti  J.  et  M.;  Stony  Creek, 
111.  (November).    The  fish  was  kept  alive  in  an  aquarium  from  November 


156  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [394 

11,  1918,  until  March  10,  1919,  when  it  was  killed,  being  then  nearly 
dead.  The  material  was  examined  on  March  15.  A  few  isolated  spores 
occurred  in  the  muscle  of  Notropis  blennius  (Homer  Park,  111.,  November). 

Vegetative  form:  In  and  between  the  muscle  bundles  of  the  myotomes. 
Size  variable.  Color  opaque  white  under  the  dissecting  microscope. 
Smallest  rounded  ameboid  forms  with  a  single  or  numerous  nuclei,  in  the 
muscle  bundle,  have  the  size  of  from  10/t  to  30n  in  greatest  diameter 
(Figs.  573  to  575).  The  largest  form  observed  was  400/x  by  120ju.  Young 
forms  without  any  diflferentiated  protoplasm,  shows  indistinct  granular 
and  reticular  structure  with  deeply  staining  spherical  or  ring-form  chro- 
matinic  granules.  The  number  of  the  nuclei  increases  with  the  growth  of 
the  body.  Larger  form  (Fig.  576),  spindle  shape,  circular  in  cross-section, 
lies  with  its  long  axis  parallel  to  the  muscle  fibres.  The  protoplasm 
vacuolated,  contained  mostly  mature  spores.  Spores  were  also  found  in 
the  state  of  diffuse  infiltration.  Polysporous. 

Spore:  Form  somewhat  variable.  Typical  form  almost  circular, 
slightly  pointed  at  the  anterior  end  (Fig.  566)  in  front  view;  spindle  shaped 
in  profile  (Figs.  569  and  570).  Sutural  ridge  marked.  Shell  uniformly 
thick,  usually  exhibiting  four  triangular  folds  on  the  surface  along  the 
posterior  margin  (Figs.  566,  568  and  571).  No  intercapsular  appendix. 
Two  pyriform  polar  capsules  are,  as  a  rule,  of  the  same  size  and  form. 
Frequent  occurrence  of  the  inequality  of  the  polar  capsules  together  with 
abnormalities  in  the  form  of  the  spore,  were  noticed  especially  among 
comparatively  young  spores.  The  granular  sporoplasm,  shows  two  spher- 
ical nuclei  when  stained.  The  iodinophilous  vacuole,  spherical  and  2fi  in 
average  diameter,  is  deeply  stained  with  Lugol's  solution.  Dimensions 
of  unstained  preserved  spores:  length  and  breadth  9  to  lO/x,  thickness 
6.5  to  7/x,  polar  capsule  6  to  7.5/x  by  2.5  to  3n. 


MYXOBOLUS  DISCREPANS  nov.  spec. 
[Figs.  597  to  601] 

Habitat:  Branchial  lamellae  of  Carpiodes  diformis;  Salt  Fork,  Urbana, 
U.S.A.  (May).  One  fish  caught,  died  (soon  after  the  capture)  two  hours 
before  being  fixed.    Length  8.5cm. 

Vegetative  form:  The  parasites  formed  numerous  cysts  on  the  branchial 
lamellae.  Cysts  slightly  yellowish  white  and  mostly  rounded  or  elongated 
along  the  lamella,  occur  in  groups,  often  occupying  the  entire  lamella. 
Infection  was  fairly  heavy.  Every  gill  arch  harbored  ten  to  twenty  cysts 
mostly  on  the  outer  surface.  Size  of  the  cyst  varies,  small  rounded  one 
500/x  in  diameter  up  to  elongated  forms  2mm.  by  0.5mm.,  the  majority 
being  from  0.5  to  1mm.  in  diameter.  The  cyst  is  surrounded  by  a  thin 
connective  tissue  layer  of  the  host.     The  protoplasm  shows  little  differen- 


395)  STUDIES  ON  MYXOSPORIDIA—KUDO  157 

tiation.  The  ectoplasm  is  a  rather  narrow  zone  around  the  entire  body 
and  the  endoplasm  is  filled  with  various  nuclei,  several  stages  of  developing 
pansporoblasts,  and  mature  spores.  Each  pansporoblast  produces  two 
spores.     Polysporous. 

Spore:  Approximately  circular  with  broad  anterior  and  more  or  less 
narrower  posterior  end  in  front  view;  broadly  fusiform  in  profile.  Shell 
uniformly  thin  with  5  to  6  markings  on  the  posterior  margin.  Two  polar 
capsules  broadly  oval  and  convergent,  fill  the  anterior  half  of  the  spore. 
A  small  triangular  intercapsular  appendix  presents.  Coiled  polar  filament 
is  fairly  visible  in  vivo.  The  spores  from  the  cysts  which  were  fixed  with 
alcohol-acetic  and  preserved  in  95  per  cent  alcohol,  showed  the  extrusion 
of  the  polar  filament  under  the  influence  of  potassium  hydrate  solution 
(35  per  cent)  even  after  a  considerable  length  of  time  as  is  shown  in  the 
following: 

Material  fixed  on  May  29.  . 

June  2;  Extrusion  took  place  in  almost  all  spores. 

June  10;  Extrusion  took  place  in  almost  all  spores. 

June  26;  Extrusion  took  place  in  almost  all  spores. 

July  28;  Extrusion  took  place  in  almost  all  spores. 

August  29;  Extrusion  took  place  in  numerous  spores. 

September  29;  Extrusion  took  place  in  about  70  per  cent  of  the  spores,  some  filaments 
being  rather  short,  and  not  fully  extended. 

October  20;  Extrusion  took  place  in  about  50  per  cent  of  the  spores,  most  filaments  being 
short,  and  not  fully  extended. 

Sporoplasm  coarsely  granular  shows  clearly  two  ring-form  nuclei  in 
fresh  preparations.  Dimensions  of  preserved  spores:  length  11.4  to 
13.5/1,  breadth  9.5  to  llju,  thickness  8.5  to  9.5ju,  polar  capsule  5.5  to  6n 
by  3.5  to  4/x,  length  of  polar  filament  50  to  55/i. 

Remarks:  The  present  species  differs  from  the  hitherto  known  species, 
Myxobolu»  lintoni  (page  138)  and  Myxobolus  orbiculatus  (page  155)  which 
are  the  nearest  to  the  present  form,  differ  from  Myxobolus  discrepans  in 
the  host,  organ  of  infection,  vegetative  form  and  form  and  structure  of 
the  spore. 

MYXOBOLUS  MESENTERICUS  nov.  spec.     , 
[Figs.  628  to  631] 

Habitat:  In  the  mesentery,  liver,  spleen  and  wall  of  stomach,  pyloric 
coecum,  intestine,  and  gall-bladder  of  Lepotnis  cyanellus;  Crystal  Lake, 
Urbana,  111.  (June  and  July).  Out  of  thirty-six  host  fish,  10  cm.  in  average 
length,  seven  were  found  to  be  infected.  In  every  case,  except  one,  the 
mesentery  was  the  main  seat  of  infection,  harboring  conspicuous  cysts. 
The  number  of  cysts  found  in  the  host  body  varied  from  three  to  seven. 
The  infected  fish  did  not  exhibit  any  recognizable  pathological  changes. 
Other  species  of  fish  caught  at  the  same  time,  were  free  from  the  infection. 


158  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [396 

Vegetative  form:  The  cysts  are  mostly  spherical  in  form,  and  are 
covered  by  a  tough  resistant  envelope  composed  of  the  connective  tissue 
of  the  host.  They  are  uniformly  white  in  color,  and  have  the  variable 
dimensions  of  from  0.5  to  1.5mm.  in  diameter.  In  section,  the  protoplasm 
shows  a  coarsely  reticulated  structure  without  distinct  differentiation. 
In  all  cysts  of  various  sizes  fully  mature  spores  were  only  observed.  The 
spore  formation  could  not  be  worked  out.     Polysporous. 

Spore:  Broadly  oval  with  a  slightly  truncated  anterior  end  in  front 
view  (Fig.  628),  lenticular  in  side  or  end  view  (Fig.  629).  No  intercapsular 
appendix  is  seen.  The  shell  is  rather  thick,  and  shows  about  eight  folds 
on  the  sutural  edge,  two  of  which  located  laterally  being  more  conspicuous 
than  others.  The  sutural  ridge  is  rather  fine.  Two  convergent  polar 
capsules  equal  in  size  occupy  the  anterior  half  of  the  spore.  The  coiled 
polar  filament  becomes  more  distinctly  visible  with  the  addition  of  Lugol's 
solution,  altho  it  is  faintly  observable  in  fresh  state.  Fresh  spores  ex- 
truded their  polar  filaments  under  the  action  of  potassium  hydrate  solu- 
tion. In  some  spores,  the  extruded  filaments  cross  each  other  near  the 
foramina.  The  preserved  spores  showed  no  extrusion  of  the  filament 
as  in  the  last  species.  The  sporoplasm  is  extremely  finely  granu- 
lated. The  iodinophilous  vacuole  is  comparatively  large.  When  stained, 
the  spore  shows  two  nuclei  in  the  sporoplasm.  Dimensions  of  fresh 
material:  length  10  to  11.5jli,  breadth  8.5  to  9.5m,  thickness  6.5ai,  polar 
capsule  4.75m  by  1.5  to  2n,  length  of  polar  filament  32  to  40^.  Average 
dimensions  of  unstained  preserved  spores:  length  9.5/*,  breadth  8m,  polar 
capsule  4.75  m  by  2m. 

Remarks :  The  habitat  and  the  structure  of  the  spores,  lead  the  writer 
to  record  the  species  as  a  new  species. 

Genus     HENNEGUYA     Thelohan 
1892        Henneguya  Thelohan  1892  :  167,  176 

1895        Henneguya  Th61ohan  1895  :  352 

The  characters  of  the  genus  are  described  on  page  59. 
Type  species :  Henneguya  psorospermica  Thelohan. 

HENNEGUYA  PSOROSPERMICA  Thelohan 
[Figs.  486,  487  and  496] 


1895 

Henneguya  psorospermica 

Thdlohan 

1895 

:ZS3 

1896 

Myxobolus  psorospermica  s. 

str.  Cohn 

1896 

261 

1899 

Henneguya  psorospermica 

typica 

Labb6 

1899 

101 

1905 

Henneguya  psorospermica 

Nufer 

1905 

:  77,  185 

1910 

Henneguya  psorospermica 

Wegener 

1910 

:  81-82 

1911 

Henneguya  psorospermica 

typica 

Auerbach 

1911 

:  5,  etc. 

Habitat:  Branchiae  of  Esox  lucius  L.  and  Ferca  fluviatilis;  France, 


397]  STUDIES  ON  MYXOSPORIDIA—KUDO  159 

Frisches  and  Kurisches  Hafif,  Pregel,  Masurische  Seen  (all  the  year  round, 
but  rarer  in  Winter)  Switzerland. 

Vegetative  form:  Thelohan's  observations  on  the  structure  of  the 
cyst,  are  as  follows:  The  surface  of  the  cyst  is  covered  by  a  la5''er,  homo- 
geneous, refringent  and  deeply  stained,  with  which  the  cyst  comes  in 
direct  contact  with  the  surrounding  epithelial  cells  of  the  host.  Inside 
of  this  layer,  there  is  a  "pseudoectoplasmic"  zone,  in  which  the  protoplasm 
is  dense  at  places,  forming  radiate  irregular  striations,  enclosing  numerous 
irregular  masses  which  are  composed  of  apparently  the  same  substance  that 
forms  the  external  layer.  Toward  the  central  portion  of  the  cyst,  there  are 
masses  of  spores  (Fig.  496). 

Cohn's  descriptions  are  as  follows:  The  purely  white  cyst  is  elliptical; 
length  1.15mm.  and  breadth  0.85mm.  The  seat  is  under  the  epidermis. 
It  is  surrounded  by  the  host  tissue  with  small,  elongated  and  scattered 
nuclei.    The  outer  layer  of  the  cyst  is  a  thin  membraneous  protoplasm. 

Wegener  writes  as  follows:  The  white  cysts  are  round  or  elliptical, 
usually  on  the  upper  end  of  the  branchial  lamella.  Size  of  larger  cysts, 
1.5  to  2mm.  long  and  1.1  to  1.5mm.  wide. 

Spore:  Elongated;  anterior  part  fusiform  and  anterior  end  blunt. 
Polar  capsules  elongated  and  parallel  to  each  other.  Coiled  polar  filament 
visible  in  fresh  conditions.  Shell  unstriated.  Dimensions:  total  length 
40)u  in  average,  largest  breadth  7/x,  length  of  polar  capsule  7  to  8/x. 

Cohn's  form  is  described  by  him  as  follows:  Spore  narrow  with  blunt 
anterior  end.  Sporoplasm  with  6  horns  (no  figure  to  explain  this  expres- 
sion!). When  kept  in  water,  sporoplasm  takes  round  form  and  becomes 
highly  refractive.  Dimensions:  length  29  to  38/x,  length  between  the  tip 
and  the  posterior  margin  of  the  cavity  (15  to  20/x)  18;u,  breadth  9  to  10/x, 
polar  capsule  (8  to  ll)u)  9/i  by  2tx,  length  of  "starren  Faden"  14/n,  length  of 
tail  14  to  18m. 

Wegener's  form  is  as  follows:  total  length  35  to  38/i,  breadth  7  to  8/*, 
length  of  the  spore  cavity  15^,  length  of  tail  15  to  20/x,  polar  capsule  8)li  by 
2  to  3)u. 

HENNEGUYA  TEXTA  (Cohn)  Labbe 

1895        Myxoholiis  textus  Cohn  1895  :  38-39 

1899        Henneguya  psorospermicatexta  Labb6  1899  :  101 

1910        Henneguya  texta  Wegener  1910  :  82-83 

Habitat:  Branchiae  of  Perca  fluviatilis  L.;  Pregel,  Frisches  and 
Kurisches  Haff  (all  the  year  round). 

Vegetative  form :  Cohn  observed  as  follows :  Cyst  distinctly  elliptical. 
Length 0.75mm.,  breadth 0.375mm.  The  cysts  surrounded  by  a  thick  layer 
of  the  host  tissue.  In  the  peripheral  portion  of  the  cyst,  the  protoplasm 
exhibits  a  network-like  structure  which  forms  a  fibrous  structure  further 
inside,  crossing  the  cyst  at  right  angles  to  the  long  axis  of  the  cyst. 


160  ILLINOIS  BIOLOGICAL  MONOGRAPHS  (398 

Wegener  writes  as  follows:  The  white  cysts  are  elongated,  1.2  to  1.8mm. 
long  and  0.5  to  0.7mm.  wide. 

Spore:  Cohn  mentions  dimensions  exactly  the  same  as  those  of 
Henneguya  psorospertnica  and  can  not  distinguish  the  two  species  by  the 
spore. 

Wegener  gives  the  following  dimensions:  length  30  to  40/i,  breadth 
7  to  8m,  length  of  the  cavity  of  spore  15  to  18/i,  length  of  tail  15  to  25^, 
polar  capsule  8/t  by  2  to  3jn. 

HENNEGUYA  MINUTA  (Cohn)  Labbe 

[Figs.  488  and  489] 

1895        Myxobolus  minutus  Cohn  1895  :  39-40 

1899        Henneguya  psorospertnica 

minuia  Labb€  1899  :  102 

Habitat:  Branchiae  of  Percafluviatilis  L.;  Frisches  Haflf,  Lesina. 

Vegetative  form:  Cohn's  description  is  as  follows:  Cysts  oval  and 
small,  difficult  to  distinguish  them  from  those  of  Henneguya  psorospermica. 
Size,  130/1  by  115/x.  The  parasite  was  met  only  once.  But  the  number  of 
the  cysts  was  far  greater  than  that  of  Henneguya  psorospertnica,  often  5  to  6 
on  one  lamella,  reaching  up  to  200  cysts  on  a  single  gillarch. 

Spore:  Cohn  gives  the  following  dimensions:  total  length  (28  to  45)u) 
about  36/i,  length  from  the  tip  to  the  end  of  cavity  (20  to  28m)  about  26m, 
breadth  10  to  11m,  thickness  8m,  polar  capsule  11  to  14m  by  2  to  Zn,  length 
of  polar  filament  42  to  45m,  length  of  tail  (8  to  17m)  12m.  Cohn  gives  a 
figure  (Fig.  489)  of  a  spore  with  two  vacuoles(?). 

HENNEGUYA  OVIPERDA  (Cohn)  Labbe 
[Figs.  490  and  491] 
1892  Weltner  1892  :  28-36 

1895        Myxobolus  oviperdus  Cohn  1895  :  40-41 

1899        Henneguya  psorospertnica  ovi- 

perda  Labbfi  1899  :  102 

1904        Hentieguya  ^psorospertnica  ovi- 

perda  Fuhnnann  1904:469-471 

19 1 1        Henneguya  psorospertnica  ovi- 

perda  Auerbach  1911  : 5-22 

1911        Henneguya  psorospertnica  ovi- 

perda  Nemeczek  1911  :  146 

Habitat:  Ovary  of  Esox  lucius  L.;  Switzerland,  Berlin,  Frisches  Haff 
(all  the  year  round),  Upsala  (May),  Austria  (December). 

Vegetative  form:  Cohn  writes  as  follows:  No  real  cyst  exists.  The 
parasite  occupies  the  ovum. 

Auerbach,  however,  mentions  the  presence  of  cysts  in  the  connective 
tissue  and  follicle  epithelium  of  the  ovary.  Dimensions,  1mm.  up  to  5  or 
6mm.  in  diameter. 


399\  STUDIES  ON  MYXOSPORJDIA—KUDO  161 

Spore:  Cohn  states  the  form  and  dimensions  are  very  much  similar 
to  those  of  H.  psorospermica. 

HENNEGUYA  LOBOSA  (Cohn)  Labbe 
[Figs.'  492  and  493] 

1895        Myxobolus  lobosus  Cohn  1895  :  42 
1899        Henneguya  psorospermica  lo- 

bosa  Labb6  1899  :  102 

1910  Henneguyai?)  lobosa  Wegener  1910  :  83 

1911  Henneguyai?)  lobosa  Auerbach  1911  :  22-25 

Habitat:  Branchiae  of  Esox  lucius  L.;  Frisches  Hafif,  Pregel,  Karlsruhe. 

Vegetative  form :  Cysts  irregular  in  shape,  size  up  to  2.5mm. 

Wegener  noticed  that  the  cyst  resembles  that  of  Myxosoma  dujardini 
with  the  dimensions  of  2.2  to  2.8mm.  by  1  to  1.1mm. 

Spore:  Cohn  gives  the  dimensions  as  follows:  total  length  30  to  40/i, 
length  from  the  tip  to  the  posterior  margin  of  cavity  11.5  to  15/i,  breadth 
5  to  6.5m,  polar  capsules  6.5  to  Sfi  by  2  to  2.5/x,  length  of  tail  22  to  ISfi. 

Wegener's  form:  oval;  length  35  to  40^1,  breadth  Sfi,  polar  capsule  6  to 
7n  by  2.5  to  3n,  length  of  the  cavity  of  spore  13  to  15^,  length  of  tail  20  to 
25/i,  the  iodinophilous  vacuole  could  not  be  detected. 

Auerbach  gave  the  following  dimensions:  total  length  30/*,  breadth 
4  to  6n,  length  of  polar  capsule  6n,  length  of  polar  filament  48  to  54/u. 

Remarks:  Wegener  and  Auerbach  did  not  observe  the  iodinophilous 
vacuole. 

HENNEGUYA  PERI-INTESTINALIS  Cepede 

1906  Henneguya  psorospermica  peri- 

intestinalis  Cfipfide  1906  :  67 

1907  Henneguya  psorospermica  peri- 

intesiinalis  C^de  1907  :  137 

1912  Henneguya  psorospermica  peri- 

intestinalis  Parisi  1912  :  295 

Habitat:  Intestine  of  Esox  lucius  L.;  Lac  du  Bourget,  Pavia.  (June). 
Vegetative  form:  Cysts. 

Spore:  Cepede  mentions  that  it  resembles  that  of  Henneguya  psoro- 
spermica. 

HENNEGUYA  MEDIA  Thelohan 
[Figs.  494  and  495] 
1890  Thelohan  1890  :  198-200 

1892        Henneguya  media  Thdohan  1892  :  177 

1894  Myxobolus  medius  Gurley  1894  :  248 

1895  Henneguya  media  Thglohan  1895:353 
1898        Henneguya  media                       Doflein  1898  :  342 

Habitat:  Renal  tubules  of  kidney  and  ovary  of  Gasterosieus  aculeatus 
and  G.  pungitius  L.;  France.    Mixed  infection  with  Sphaerospora  elegans. 


162  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [400 

Vegetative  form:  Rounded  or  elongated.  In  larger  individuals,  clear 
differentiation  of  protoplasm.    Monosporous  (?)  and  polysporous. 

Spore:  Fusiform.  Shell  striated.  A  vacuole  in  sporoplasm.  Dimen- 
sions: length  20  to  24ju,  breadth  5  to  6/t,  polar  capsules  4  to  5/x.    Tail  short. 

HENNEGUYA  BREVIS  Thelohan 
1854  Lieberkuhn         1854  :  357 

1892  Henneguya  brevis  Th61ohan  1892  :  177 
1895        Henneguya  brevis                      TWlohan             1895:354 

Habitat:  Similar  to  H.  media  Thelohan. 
Vegetative  form:  Undescribed. 

Spore:  Fusiform  with  short  tail.  Dimensions:  length  14  to  ISju, 
breadth  5  to  6/1,  polar  capsules  1.4  to  5jU,  tail  4  to  5)u  long. 

HENNEGUYA  SCHIZURA  (Gurley)  Labb6 

[Figs.  497  to  499] 

1841  MuUer  1841  :  477-478 

1893  Myxobolus  schizurus  Gurley  1893  :  417 

1894  Myxobolus  schizurus  Gurley  1894:255 
1899        Henneguya  schizura                   Labb6  1899  :  102-103 

Habitat:  In  cellular  tissue  of  the  eye  muscles,  in  that  of  the  sclerotic, 
and  in  that  between  the  sclerotic  and  choroid  of  Esox  lucius  L.;  Germany, 
U.  S.  A. 

Vegetative  form:  Cysts  white;  membrane  delicate;  0.44  to  1.09mm.  in 
diameter. 

Spore:  Oval.  Dimensions:  length  12/x,  breadth  6/i,  thickness  one- 
half  the  breadth,  tail  3  to  4  times  length  of  the  body. 

HENNEGUYA  CREPLINI  (Gurley)  Labbe 


[Figs. 

500  to  503] 

1842 

Creplin 

1842 

:  61-63 

1894 

Myxobolus  creplini 

Gurley 

1894 

:  248-249 

1899 

Henneguya  creplini 

Labb€ 

1899 

:103 

1910 

Henneguya  creplini 

Wegener 

1910; 

:84 

Habitat:  Branchiae  of  Acerina  cernua  L.;  Pregel  (March),  Frisches 
and  Kurisches  Haff. 

Vegetative  form:  Wegener  describes  as  follows:  Cysts,  usually  elon- 
gated oval  and  are  located  at  the  end  of  branchial  lamella.  Color  white. 
Size  1  to  1.1mm.  by  0.5mm.  During  winter,  the  cyst  has  only  pansporo- 
blasts, but  no  fully  grown  spores. 

Spore:  Creplin  writes  as  follows:  Elongated  elliptical.  Length  1/120'", 
breadth  1/360'",  tail  about  as  long  as  or  a  little  longer  than  the  body. 

Wegener's  form:  elongated  spindle  shape;  length  20At,  breadth  8  to 
9iJL,  polar  capsule  8/i  by  2  to  3/x  (parallel  to  each  other). 


4011  STUDIES  ON  MYXOSPORJDJA—KUDO  163 

Remarks:  Wegener  thinks  that  the  present  species  and  Henneguya 
acerinae  Schroder,  are  one  and  the  same  species,  and  that  the  differences 
between  the  dimensions  are  due  to  the  miscalculation  of  measurement  in 
lines  given  by  Creplin  on  the  part  of  Gurley  and  Labbe. 


HENNEGUYA  LINEARIS  (Gurley)  Labbe 
[Fig.  504] 

1841  MuUer  1841  :  489 

1893  Myxobolus  linearis  (part)  Gurley  1893  :  417 

1894  Myxobolus  linearis  Gurley  1894  :  255 
1899  Henneguya  linearis  Labb6  1899  :  103 

Habitat:  Membrane  lining  branchial  cavity  of  Pimelodus  sebae  Cuv. 
et  Val.,  branchiae  of  Platystoma  fasciatum  L.;  South  American  rivers. 
Vegetative  form:  Not  described. 
Spore:  Very  narrow.    Length  3  to  4  times  breadth. 


HENNEGUYA  GURLEYI  Kudo 

[Fig.  505] 

1893  Myxobolus  linearis  ipaxt)  Gurley  1893:417 

1894  Myxobolus  ci.  linearis  Gurley  1894:253-254 
1899        Henneguya  linearis  v&T.             Labb6                   1899  :  103 

Habitat:  Base  of  spines  of  the  second  dorsal  fin  of  Ameiurus  melas 
Raf.;  Iowa  (Storm  Lake)  (August). 

Vegetative  form:  Spherical  cysts,  1mm.  in  diameter. 

Spore:  Lanceolate.  Dimensions:  length  of  the  body  19/i,  width 
5  to  6/i,  thickness  about  3)it. 

Remarks:  The  species  is  most  probably  different  from  Henneguya 
linearis  judging  from  the  difference  in  the  form  and  structure  of  spores, 
the  seat  of  infection,  and  host  species.  Hence,  it  is  recorded  here  as  an 
independent  species. 


HENNEGUYA  STRONGYLURA  (Gurley)  Labb6 
[Fig.  506] 
1841  MuUer  1841  :  480 

1894        Myxobolus  strongylurus  Gurley  1894:249 

1899        Henneguya  strongylura  Labb6  1899:103 

Habitat:  Skin  of  cephalic  region  of  Synodontis  schall  Bl.  Schn.;  Nile. 
Vegetative  form:  Cysts  over  2.18mm.  in  diameter. 
Spore:  Dimensions:  length  of  the  body  9/i,  breadth  5.4/1.    Tail  always 
undivided.    Two  polar  capsules  of  equal  size. 


IW  ILUNOJS  BIOLOGICAL  MONOGRAPHS  (182 

HENNEGUYA  MONURA  (Gurley)  Labbe 
[Fig.  507] 

1880  Ryder  1880  :  211-212 

1893  Myxobolus  monurus  Guriey  1893  :  416 

1894  Myxobolus  monurus  Guiley  1894  :  249-250 
1899  Eentuguya  monura  Labb6  1899  :  103 

Habitat:  Subcutaneous  intermuscular  tissue  of  Aphredoderus  sayanus 
Gill.;  New  Jersey  (Woodbury). 

Vegetative  form:  Cysts,  lenticular,  large,  white,  opaque  and  numerous 
(20).     Membrane  thin. 

Spore:  Lenticular  or  slightly  obovate.  Tail  2  to  3  times  longer  than 
the  body. 

HENNEGUYA  KOLESNIKOVI  (Gurley)  Labbe 
[Fig.  508] 

1886  Kolesnikov  1886  :  242-248 

1894        Myxobolus  koksnikovi  Gurley  1894  :  256-257    • 

1898  Myxobolus  bicaudatus  (part)     Zschokke  1898  :  602-604,  646- 

655,  699-703 

1899  Hmneguya  koUsnikovi  Labh€  1899  :  103-104 

Habitat:  Interstitial  connective  tissue  of  the  thoracic  and  intercostal 
muscles  of  Coregonus  lavaretus  L.;  Russia. 

Vegetative  form:  Cysts  numerous  (80),  spherical  or  oval;  length  10 
to  30mm.,  breadth  7  to  20mm. 

Spore:  Oval  with  a  pointed  anterior  end.  Tail  three  times  longer  than 
the  body. 

Remarks:  Zschokke  thinks  the  present  species  is  identical  with  Henne- 
guya  zschokkei.  But  the  evidence  is  not  clear  enough  to  bring  one  to 
agree  with  him  due  to  the  incomplete  description  of  the  present  species. 


HENNEGUYA  MACRURA  (Gurley)  Thelohan 
[Figs.  509  to  512] 

1893  Evennann  1893  :  76 

1894  Myxobolus  macrurus                  Gurley  1894  :  250-253 

1895  Henneguya  macrura                  Thdohan  18^5  :  354 

Habitat:  Subcutaneous  connective  tissue  of  head  of  Hybognaihus 
nuchalis  Ag.;  Neches  River,  Texas  (November,  temperature  of  water 
9*'.4C.)    Of  frequent  occurrence. 

Vegetative  form:  Cysts,  elongated  6mm.  by  2mm.  or  less. 

Spore:  Rounded  oblong.  Dimensions:  length  10  to  11m,  breadth 
6  to  8/x,  thickness  4^.    Shell- valves  unequally  convex.    Tail  30  to  40^. 


403]  STUDIES  ON  MYXOSPORIDJA—KUDO  165 

HENNEGUYA  ZSCHOKKEI  (Gurley)  Doflein 
[Fig.  513] 


1884 

Zschokke 

1884  : 

:  234-235 

1894 

Myxoholus  (?)  zschokkei 

Gurley 

1894  : 

244 

1898 

Myxobolus  bicaudatus  (part] 

1  Zschokke 

1898 

:  602-607,  646- 
655,  699-703 

1898 

Myxoholus  bicaudatus 

Zschokke 

1898a 

:  213-214 

1901 

Henneguya  zschokkei 

Doflein 

1901  ; 

:202 

1904 

Henneguya  zschokkei 

Hofer 

1904; 

:56 

1905 

Henneguya  zschokkei 

Nufer 

1905 

:  77,  185 

Habitat:  Subcutaneous  and  superficial  intermuscular  tissue  of  Core- 
gonusfera,  C.  schinzii  Fatio,  C.  hiemalis  Jur.  and  muscular  tissue  and  bran- 
chia  of  C.  wartmanni  nobilis  and  C.  exiguus  albellus;  Neuch^teler  See,  Zurich 
See,  Genfer-see,  Thuner-see,  Vierwaldstatter-see. 

Vegetative  form:  Zscholclce  writes  as  follows:  Cysts  rounded  or  oval 
surrounded  by  a  compact  membrane  with  many  nuclei.  The  largest 
32mm.  by  16mm.    Protoplasm  granular.     Polysporous. 

Spore :  Rounded  oval  in  front  view ;  broad  elliptical  in  side  view.  Anter- 
ior end  rounded^  posterior  end  tapering,  forming  tail.  Sutural  ridge 
distinct.  Tail  is  either  bifurcated  along  the  entire  length  or  a  single  form, 
no  intermediate  form  being  observed.  Dimensions:  total  length  5Sfx, 
length  of  the  body  10)u,  breadth  7/i,  length  of  tail  4  to  5  times  the  length 
of  the  spore-body,  length  of  polar  filament  6  to  10  times  that  of  the  body 
of  the  spore. 

Remarks:  Zschokke  thinks  that  Henneguya  kolesnikovi,  H.  zschokkei 
and  H.  sp.  Gurley  are  one  and  the  same  species,  for  which  he  proposed 
the  name  Myxobolus  bicaudatus. 

HENNEGUYA  sp.  (Gurley)  Labbe 
[Fig.  514] 
1886  Benecke  1886  :  211 

1894        Myxoholus  sp.  inc.  Gurley  1894  :  244 

1899        Henneguya  sp.  Labbe  1899  :  104 

1904        Henneguya  sp.  Hofer  1904  :  51 

Habitat:  Integument  (?)  of  Leuciscus  rutilus  L.    The  parasites  formed 
boil-like  enlargement  in  the  skin. 
Vegetative  form:  Not  described. 
Spote:  Not  described. 


HENNEGUYA  sp. 

(Gurley) 

Labbe 

1874 

Claparede 

1874 

:114 

1894 

Myxoholus  sp.  inc. 

Gurley 

1894; 

;253 

1898 

Myxobolus  bicaudatus  (part) 

Zschokke 

1898 

:  602-607,  646- 
655,  699-703 

1899 

Henneguya  sp. 

Labbe 

1899; 

:104 

Habitat:  Branchial-arches  of  Coregonus /era;  Genfer-see. 


166  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [404 

Vegetative  form:  One  cyst,  1mm.  in  diameter. 
Spore:  Tail  short.  Zschokke  quotes:  length  8  to  lO/i. 
Remarks:  According  to  Zschokke,  this  species  is  identical  with  H. 
zschokkei. 

HENNEGUYA  TENUIS  Vaney  et  Conte 
[Fig.  515] 
1901        Hmneguya  tenuis         Vaney  and  Conte  1901  :  103-106 

Habitat:  Connective  tissue  of  alimentary  tract  of  Acerina  cernua  L.; 
Lyon  (February). 

Vegetative  form:  Numerous  cysts  particularly  in  the  pyloric  coecum. 
Usually  spherical.    Size:  30  to  150/*  in  diameter. 

Spore:  Oval  and  small.  Tail  short.  Two  polar  capsules  at  the  anterior 
end.  Sporoplasm  with  a  nucleus,  rod-shaped,  with  somewhat  enlarged 
ends  which  is  located  at  right  angles  to  the  longitudinal  axis.  lodinophilous 
vacuole  could  not  be  traced.    Dimensions:  length  4/i,  breadth  2/i. 

HENNEGUYA  NUSSLINI  Schuberg  et  Schroder 

[Figs.  516  and  517] 

1905        Hettneguya  nUsdini         Schubetg  and  Schroder         1905  :  56-59 

Habitat:  Subcutaneous  connective  tissue  at  the  base  of  dorsal  fin  of 
Truitafario  L.;  Gutach. 

Vegetative  form:  Trophozoites  form  cysts  (2  cysts  found).  Cysts 
lenticular,  1.5  to  2mm.,  surrounded  by  many  concentric  layers  of  fibrous 
connective  tissue.    Cysts  containing  only  mature  spores. 

Spore:  Broad  oval  form,  flattened.  Anterior  end  rounded.  Tail  at 
the  posterior  end.  Shell  somewhat  thick,  often  shows  sutural  ridge.  Tail 
filaments  two.  A  "dark  part"  which  in  side-view  is  of  triangular  form, 
runs  into  the  tail.  Sporoplasm,  occupying  the  posterior  half  of  the  spore, 
projects  a  narrow  portion  between  the  polar  capsules  beyond  the  middle  of 
the  capsules.  Sporoplasm,  uniformly  granular,  contains  an  iodinophilous 
vacuole  and  one,  sometimes  two  nuclei  connected  by  nuclear  bridge. 
Polar  capsules,  pyriform,  opening  independently.  Coiled  polar  filament 
observable,  coiled  6  to  7  times.  Dimensions:  length  excluding  tail  12^, 
length  with  tail  32/x,  breadth  8  to  9/x,  polar  capsules  Sn  by  3ai,  length  of 
polar  filament  4  to  5  times  longer  than  that  of  spore  excluding  tail  (48  to 
60m). 

HENNEGUYA  L£GERI  Cepede 
[Figs.  518  to  523] 

1905  Hameguya  Ugen  Cepede  1905  :  905-913 

1906  Heitneguya  Ugeri  Cepede  1906  :  66 
1913        Hermeguya  Ugen                      Cepede                1913  :  302-305 

Habitat:  Urinar\'  bladder  of  Cohitis  barbaittla  L.;  Isere  (January). 


405]  STUDIES  ON  MYXOSPORIDIA—KUDO  167 

Vegetative  form:  Young  trophozoites  subcircular,  irregularly  elliptical 
or  elongated  with  distinct  differentiation  of  protoplasm  into  ectoplasm 
and  endoplasm.  Plasmotomic  multiplication  takes  place  during  winter 
months,  when  no  spore  is  formed. 

Spore:  Oval  with  short  tail,  mostly  bifurcated  at  the  free  end.  The 
anterior  end  is  more  rounded,  occasionally  acuminated.  Two  polar  capsules 
of  equal  size.  Coiled  polar  filament  distinct  in  vivo.  Sporoplasm  granular, 
contains  two  nuclei  and  a  vacuole.  The  spore  often  shrinks  in  fresh  con- 
ditions, probably  owing  to  the  poorly  developed  thin  valves.  Dimensions 
of  spores  mounted  in  balsam:  length  variable.  Examples:  Total  length 
22.5m,  tail  8.5/i;  total  length  19.5pt,  tail  8/i;  length  of  main  part  8.5/^,  breadth 
(comparatively  constant)  6/*. 

HENNEGUYA  ACERINAE  Schroder 
[Figs.  525  and  526] 

1906        Henneguya  acerinae  Schroder  1906  ;  186-196 

1910  Henneguya  creplini  Wegener  1910  :  84 

1911  Henneguya  acerinae  Nemeczek  1911  :  155 

Habitat:  Branchiae  of  Acerina  cernua  L.,  Aspro  zingel  Cuv.,  Lucioperca 
lucioperca  L.  and  L.  sandra  Cuv.  (?) ;  Heidelberg  (Necker),  Apatin,  Komitat 
Baco-Bodrog,  Hungary  (May). 

Vegetative  form:  Schroder  describes  as  follows:  Rounded  or  spherical 
cysts  in  the  connective  tissue  of  branchial  lamella.  Full-grown  cysts  up  to 
300/1  in  diameter.  Protoplasm  is  differentiated  into  ectoplasm  and  endo- 
plasm. Ectoplasm  shows  fine  radial  striations.  Endoplasm  granular, 
contains  many  nuclei,  especially  lying  in  the  middle  portion.  Well  devel- 
oped cyst,  containing  only  spores,  is  surrounded  by  a  membrane.  On  the 
surface  of  the  ectoplasm,  numerous  edge-like  elevations,  branched  and 
joining  together,  were  recognized.    Polysporous. 

Nemeczek  observed  the  largest  cyst,  spherical  and  600//  in  diameter.    . 

Spore:  Pyriform  in  front  view;  flattened.  The  anterior  end  is  more  or 
less  blunt.  Shell  uniformly  thin.  Sutural  edge  slightly  enlarged.  Sporo- 
plasm finely  granular,  contains  an  iodinophilous  vacuole  and  two  nuclei. 
Polar  capsules  approximated  closely,  each  having  an  independent  opening. 
Dimensions:  length  20  to  22)li,  breadth  8  to  9/i,  thickness  6  to  7/x,  length  of 
tail  50  to  60m,  polar  capsules  lO/x  by  2  to  3)u,  length  of  polar  filament  80  to 
90/ii  (water  and  nitric  acid). 

Nemeczek's  form  is  as  follows: 

The  tail  is  bifurcated  along  its  entire  length.  In  one  case  (May,  1909), 
however,  all  the  spores  had  no  bifurcated  tail,  while  the  polar  capsules  were 
of  unequal  size.  Dimensions  in  fresh  state:  total  length  37.6  to  41. 8/i, 
length,  excluding  tail  12.6  to  16.8m,  breadth  4.5/x,  length  of  polar  capsule 
6.3  to  8.4jLi,  length  of  polar  filament  67/x,  length  of  tail  25m. 


168  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [406 

Nemeczek  observed  two  more  different  (?)  forms.  One  form  found  in 
Lucioperca  sandra,  tho  the  size  differs  from  the  dimensions  given  by 
Schroder,  is  thought  to  be  identical  with  the  present  species.  Another 
form  in  the  branchiae  of  As  pro  zingel,  which  is  also  to  be  one  and  the  same 
species  with  the  present  species  has  the  following  dimensions :  total  length 
35)u,  length  of  spore  excluding  tail  15)u,  breadth  Sn,  length  of  polar  capsule 
6fx,  length  of  tail  20/x. 


HENNEGUYA  GIGANTEA  Nemeczek 
[Figs.  527  to  535] 

1911        Henneguya  giganlea  Nemeczek  1911:146-154 

1914        Henneguya  gigantea  Georg6vitch         1914  :  387-409 

Habitat:  Branchiae  of  Lucioperca  sandra  Cuv.;  Apatin,  Komitat 
Bacs-Bodrog,  Hungary,  Petrograd.  Nemeczek  mentions  that  the  infection 
takes  place  only  among  young  fish. 

Vegetative  form:  Cysts  numerous  and  of  conspicuous  size  in  the  free 
end  of  branchial  lamella.  In  average,  each  gill-arch  has  about  100  cysts 
which  are  of  creamy  color.  Young  cysts  400  to  450/i  in  diameter.  They 
gradually  begin  to  increase  the  size,  from  autumn  until  toward  the  end  of 
spring,  during  which  period,  the  contents  remaining  in  the  stages  of  pan- 
sporoblasts formation.  Older  cysts  rounded  spindle  shape  with  the  length 
of  4  to  7mm.  and  the  breadth  of  2  to  3mm.  The  connective  tissue  and 
epithelial  cell  layers  form  the  cyst  membrane.  The  connective  tissue 
either  simply  surrounds  the  parasite  or  branches  in  the  surface  of  the  para- 
site, increasing  in  thickness  and  forming  more  or  less  enclosed  chambers 
of  the  parasite.  The  membrane  of  the  cyst  which  contains  mature  spores 
is  usually  very  thin.  Throughout  the  growth  of  the  cyst,  "chromatoid 
body"  is  seen  in  the  endoplasm,  which  appears  first  as  a  filiform  struc- 
ture,, stained  deeply  with  nuclear  stain.  Later  they  gather  together  and 
form  a  compact  body,  situated  excentrically.  Fine  branches  from  it 
become  directed  toward  the  surface  of  the  body,  anastomosing  each  other 
so  that  a  network  is  formed  on  the  surface  of  the  cyst.  The  latter  develops 
small  ovoidal  or  columnal  bodies  (1.2/u  long  and  about  In  wide),  which 
are  arranged  radially  and  densely.  The  number  and  quantity  of  these  bod- 
ies increase  in  proportion  to  the  number  of  propagative  nuclei  and  they 
begin  to  disappear,  first  in  the  central  portion,  then  in  the  periphery, 
so  that  in  fully  grown  cysts  (in  summer  months)  these  chromatoidal  bodies 
are  more  rudimentary.  Differentiated  protoplasm  is  only  recognized  in 
young  individuals,  in  which  case  ectoplasm  is  homogeneous  and  endoplasm 
reticular.     Polysporous. 

Spore:  Nemeczek  gives  the  following  accounts. 


407]  STUDIES  ON  MYXOSPORJDIA—KUDO  169 

Spindle  shape,  with  truncate  anterior  end  and  very  long  thread  like 
tail  at  the  posterior  end.  The  tail  seems  split  into  two  at  about  the  middle 
part  of  its  length.  Gentian  violet  stains  the  tail  so  intensively  that  its 
entire  length  could  easily  be  made  out.  Dimensions:  total  length  87.5  to 
IIO.Sm,  length  of  the  body  10.5^,  breadth  5/i,  length  of  tail  77  to  lOO/ii, 
length  of  polar  capsule  5/i,  length  of  polar  filament  70/i  (pressure  or  dessica- 
tion  followed  by  immersion  in  water). 

Georgevitch's  form:  length  excluding  tail  15jli,  breadth  6/i,  length  of 
tail  75^1,  length  of  polar  capsule  6/*,  length  of  polar  filament  75/*,  diameter  of 
the  iodinophilous  vacuole  4/i. 

Remarks:  Nemeczek  mentions  that  from  October  on,  cysts  had  no 
spores,  only  containing  propagative  cells.  The  velocity  of  the  development 
of  spores  depends  upon  the  temerature  of  water. 

Georgevitch  worked  out  the  spore  formation  of  the  species  and  observed 
that  the  binucleated  sporeplasm  emerged  from  the  posterior  end  of  the 
spore. 

HENNEGUYA  (?)  sp.  Nemeczek 
[Figs.  536  to  539], 

1911  Henneguya  sp.  Nemeczek  1911  :  1S7-1S9 

Habitat:  Branchiae  of  Abramis  bratna;  Komorn,  Komitat  Komorn, 
Hungary  (March). 

Vegetative  form:  Cysts  in  the  branchiae. 

Spore:  Besides  normal  spores  of  Myxobolus  rotundus  (page  149),  spores 
of  Henneguya  type  in  small  number  were  found.  The  anterior  part  of 
these  spores  resembles  that  of  the  species  mentioned  above,  while  the 
breadth  is  much  smaller  (8/i)  than  the  latter.  Majority  of  spores  have  a 
thread  like  tail,  10  to  15/i  long,  which  was  often  bifurcated.  An  iodinophil- 
ous vacuole  was  fairly  marked. 

Remarks :  It  is  placed  here  as  a  species  of  Henneguya  by  reason  of  the 
bifurcate  tail. 

HENNEGUYA  GASTEROSTEI  Parisi 
[Figs.  540  to  543] 

1912  Henneguya  gasterostei  Parisi  1912  :  296-297 

Habitat:  Kidney  of  Gasterosteus  aculeatus  L.;  Lago  di  Garda  (Feb- 
ruary). 

Vegetative  form:  Rounded  or  oval,  usually  with  two,  but  rarely  with 
four  spores.  Ectoplasm  thin  and  hyaline.  Endoplasm  contains  numerous 
granules,  most  probably  of  fatty  nature  and  decreasing  in  number  as 
spores  grow.  Free  full-grown  spores  were  seen  abundantly  in  the  connec- 
tive tissue  of  renal  tubules,  glomeruli,  etc.    Disporous  and  polysporous. 


170  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [¥» 

Spore:  Oval  with  slightly  attenuated  anterior  end;  posterior  end 
tapering  into  tails,  which  end  in  one  point  or  bifurcated;  asymmetrical  in 
shape,  one  valve  is  more  curved  than  the  other.  This  asymmetry  of  the 
sheU-valves  in  profile  enables  the  present  species  to  be  distinguished  from 
other  species.  Shell  striated  longitudinally.  Two  polar  capsules  pyriform 
and  well  developed,  reaching  to  the  middle  of  the  spore.  Sporoplasm  with 
a  round  iodinophilous  vacuole.  Dimensions:  total  length  38  to  48;*, 
length  of  the  cavity  of  the  spore  IS^t,  breadth  6  to  7.5/i,  polar  capsules  7.5 
to  9jtt  by  3  to  3.5/i,  length  of  polar  filament  50/*. 

HENNEGUYA  NEAPOLITANA  Parisi 

[Figs.  544  and  545] 

1912        Eenneguya  neapolUana  Parisi  1912  :  297-298 

Habitat:  Connective  tissue  of  the  renal  tubule  of  kidney  of  Box  sal  pa 
C.  et  v.;  Napoli  (August). 

Vegetative  form:  Small  cyst  (40  to  50/t  in  diameter)  surrounded  by 
thin  membrane,  containing  a  number  of  spores,  numerous  pigment  granules 
and  coarse  yellowish  globules. 

Spore:  Oval,  slightly  flattened.  Anterior  end  rounded  when  seen  from 
the  front,  but  attenuated  in  profile.  Shell  tapering  into  a  long  fine  tail 
posteriorly.  The  fine  distal  portion  of  the  tail  wraps  around  the  thicker 
part.  Two  polar  capsules,  pyriform,  occupying  the  anterior  half  of  the 
cavity  of  the  spore,  cross  each  other  when  seen  from  the  front.  Sporoplasm 
finely  granular  with  two  nuclei,  the  iodinophilous  vacuole  being  hardly 
visible.  Dimensions:  total  length  50  to  60ix,  length  of  the  cavity  of  spore 
8.5  to  9.5/4,  breadth  8.5  to  9.5/i,  internal  breadth  6.3  to  7/i,  thickness  8/*, 
polar  capsules  4.7  to  5.5/*  by  3/t.  « 

HENNEGUYA  WISCONSINENSIS  Mavor  et  Strasser 

[Figs.  558  and  559] 

1916       Eenneguya  wisconsinensis         Mavor  et  Strasser  1916  :  676-682 

Habitat:  Urinary  bladder  of  Perca  flavescens;  Lake  Mendota,  Wiscon- 
sin (April). 

Vegetative  form:  Trophozoites  are  usually  elongated  and  have  the 
general  form  and  shape  of  a  limax  ameba.  It  may  reach  a  size  of  300/i  by 
70/i.  Clear  differentiation  of  ectoplasm  and  endoplasm.  Pseudopodia 
lobose.    Two  spores  are  formed  in  each  pansporoblast.    Polysporous. 

Spore:  Ovoid,  bilaterally  symmetrical,  and  have  a  bifurcated  caudal 
process.  Two  polar  capsules  at  anterior  end.  Coiled  polar  filament  visible 
in  vivo  (5  windings).  Dimensions:  length  excluding  tail  11.5/i,  breadth 
7/i,  tail  9.6;t,  polar  capsules  3.5ft  by  2.5/i,  length  of  filament  33/i. 


4091  STUDIES  ON  MYXOSPORIDIA— KUDO  171 

HENNEGUYA  BRACHYURA  Ward 
[Figs.  650  to  653] 
1919        Henneguya  brachyura         Ward  1919  :  57 

Habitat:  In  the  cartilageous  fin  ray  of  the  caudal  fin  of  Notropis 
anogenus]  Put-in-Bay,  Lake  Erie  (August).  The  species  was  found 
encysted  in  the  same  fish  which  was  heavily  infected  by  Myxobolus  aureatus. 

Vegetative  form:  Cysts  rounded  with  slightly  irregular  contour  im- 
bedded in  the  fin  ray.  The  size  varies  from  160^  in  diameter  up  to  360/*  by 
240/x.  No  particular  cyst  membrane  could  be  recognized.  The  differen- 
tiation of  the  protoplasm  into  ectoplasm  and  endoplasm  is  distinct.  The 
ectoplasm  covering  the  entire  surface  of  the  parasite  as  a  layer  4  to  6/it 
thick,  shows  structure  of  a  very  finely  granular  nature.  The  endoplasm 
coarsely  alveolar,  is  filled  with  mature  spores  in  the  central  portion,  while 
numerous  nuclei  and  young  spores  in  various  developmental  stages  are 
present  at  the  peripheral  portion.     Polysporous. 

Spore:  Rounded  oval  in  front  view;  spindle  shape  with  symmetrically 
built  valves  in  profile.  Shell  rather  thick.  Sutural  ridge  fairly  well 
marked;  sutural  edge  exhibiting  a  variable  number  of  folds  (8  to  10). 
Two  pyriform  polar  capsules  are  usually  of  the  same  size  and  form.  The 
tail  is  a  single  process,  usually  more  or  less  bent  or  irregularly  curved, 
very  rarely  being  straight.  In  general,  it  is  sinuous  with  two  or  three 
shallow  curves  and  is  rather  short,  tapering  gradually  to  a  point.  In 
young  spores  which  are  less  deeply  stained  by  any  stain,  various  develop- 
mental stages  of  the  tail  are  reasily  recognized.  Giemsa  solution  stains 
the  shell  proper  in  clear  blue,  while  the  tail  takes  on  a  beautiful  pink 
color,  a  distinct  difference  in  affinity  for  dyes  between  the  material  in  the 
tail  and  the  shell.  It  seems  probable  that  the  tail  of  this  type  is  entirely 
different  in  its  development  from  that  of  the  ordinary  bifurcated  type. 
Dimensions  in  section:  length  10  to  ll.5fi,  breadth  8  to  8.75)u,  thickness 
4  to  5fi,  polar  capsules  3  to  4/x  by  2n,  length  of  the  tail  up  to  17/*. 

HENNEGUYA  SALMINICOLA  Ward 
[Figs.  654  to  656] 
1914      IHenneguya  zschokkei  Zschokke  and  Heite    1914  :  200-201 

1919        Henneguya  salminicola       Ward  1919  :  59 

Habitat:  In  the  sub-dermal  tissue  of  Onchorhynchus  keta  and  0.  kisutch 
(Zschokke  and  Heitz,  Kamtschatka)  and  in  the  connective  tissue  in  body 
muscles  of  Oncorhynchus  keta,  Stickeen  River,  Alaska  (Ward,  September). 
The  last  named  author  undertook  a  careful  examination  of  a  part  of  the 
infected  tissue  preserved  in  formol.  The  species  forms  conspicuous  C)^ts 
in  the  muscle  from  the  sub-peritoneal  to  the  sub-dermal  connective  tissue, 
tho  all  are  sub-peritoneal  in  position. 


172  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [410 

Vegetative  form:  Ward  describes  as  follows:  The  whitish  opaque  cysts 
are  pjniform,  and  fairly  uniform  in  size  (3  to  6mm.  in  diameter).  The 
cyst  is  covered  by  numerous  layers  of  connective  tissue  which  form  a 
tough  membrane  around  the  parasite.  The  cyst  contains  young  spores 
in  various  stages  of  development,  which  showed  that  two  spores  are  formed 
in  one  pansporoblast,  and  mature  spores  thickly  massed  together  in  the 
central  area.     Polysporous. 

Spore:  Oval  with  rounded  anterior  and  more  or  less  attenuated  poster- 
ior ends;  elliptical  in  profile  with  attenuated  anterior  end.  Shell  smooth. 
Sutural  edge  exhibits  folds  variable  in  number  (usually  6  to  7).  Tail 
double,  composed  of  two  fine  and  equal  halves  which  are  the  prolongation 
of  the  shell  valves.  The  processes  usually  run  roughly  parallel  to  each 
other.  Two  pyriform  polar  capsules  are  of  slightly  difiFerent  dimensions. 
Coiled  polar  filament  is  indistinct  in  preserved  unstained  specimens. 
Sporoplasm  finely  granular,  shows  a  large  iodinophilous  vacuole.  Dimen- 
sions of  stained  and  mounted  spores:  total  length  47m  (42.75  to  52.44/i), 
length  of  the  main  part  Un  (11.97  to  14.25>x),  breadth  8^  (7.12  to  8.43m), 
thickness  4.78At,  length  of  tail  35/*  (30.78  to  38.  19m),  polar  capsule  3.70  to 
4.55m  by  1.59  to  2.85m. 

Remarks:  Zschokke  and  Heitz  (1914)  observed  a  species  from  Kam- 
tschatka,  which  they  thought  to  be  identical  with  Henneguya  zschokkei 
(page  165).  The  writer  is  inclined  to  think  that  the  species  is  identical 
with  the  species  just  described  from  Alaska. 

HENNEGUYA  MIYAIRII  nov.  spec. 
[Fig.  524] 
1909        Henneguya  sp.  *  Miyairi  1909  :  127-129      < 

Habitat:  Subcutaneous  tissue  of  head  of  Carassius  auratus  L.;  Fukuoka 
(Nippon). 

Vegetative  form:  Trophozoites  form  cysts  and  are  also  found  in  the 
condition  of  diffuse  infiltration  around  the  cysts.  Cyst-membrane  fibrous 
and  thin.  Ectoplasm  and  endoplasm  fairly  well  differentiated,  though 
the  border  line  is  not  sharply  marked.  At  the  periphery  of  endoplasm, 
pansporoblasts  with  7  to  12  nuclei  are  present  (Two  spores  are  formed  in 
each  pansporoblast?).    Polysporous. 

Spore:  Oval,  with  broadly  rounded  anterior  and  slightly  elongated 
posterior  ends,  the  latter  ending  in  long  and  fine  tails.  Two  polar  capsules 
at  the  anterior  portion,  are  pyriform,  small  and  convergent.  Sporoplasm 
with  an  iodinophilous  vacuole.  Dimensions:  length  12m,  breadth  8m, 
length  of  the  tails  10  to  30m,  length  of  polar  filaments  23  to  40m. 

Remarks:  As  the  description  gives  the  details  by  which  the  species 
can  be  distinguished  from  other  species,  the  writer  estabUshes  it  on  an 
independent  basis. 


4111  STUDIES  ON  MYXOSPORIDIA—KUDO  173 

HENNEGUYA  MICTOSPORA  nov.  spec. 
[Figs.  546  to  557] 

Habitat:  Urinary  bladder  of  Lepomis  cyanellus  Raf.,  L.  humilis  Gir. 
and  Micropterus  salmoides  Lac;  Stony  Creek,  111.  (November). 

In  one  out  of  three  (6.5  to  8cm.  long)  of  the  first,  in  one  out  of  two 
(7  and  9.5cm.  long)"  of  the  second  and  in  one  of  the  third  species,  examined 
in  the  middle  of  November,  was  found  the  present  form.  None  showed  a 
heavy  infection,  a  number  of  scattered  trophozoites  and  spores  being 
observed.    The  host  did  not  show  any  pathological  change. 

Vegetative  form :  Polymorphous.  Generally  rounded  or  elongated  oval. 
In  small  monosporous  and  disporous  forms,  the  tail  of  the  spores  developed 
inside,  is  extruded  from  the  body,  so  that  these  trophozoites  show  long 
processes  (Figs.  546, 553, 555).  Pseudopodia  lobose,  and  extruded  from  the 
entire  surface  of  the  body  (Fig.  547),  tho  sometimes  they  are  well  formed 
at  one  end  of  the  body.  Protoplasm  is  differentiated  distinctly  into  ecto- 
plasm and  endoplasm.  Ectoplasm  is  homogeneous  and  hyaline,  forming 
the  outer  layer.  Endoplasm  is  of  reticular  structure.  The  body  is  colorless, 
often  yellowish,  when  the  endoplasm  is  loaded  with  numerous  yellowish 
coarse  granules.  The  size  varies  from  6  or  7/i  up  to  60)Lt.  In  a  rounded  form 
of  38/i  in  longest  diameter,  five  pansporoblasts,  each  developing  two  spores 
and  many  nuclei  were  observed.  In  another  oval  form  of  45^  by  60/*  in 
size,  numerous  nuclei  were  stained,  showing  that  no  development  of 
pansporoblast  has  yet  taken  place.  Disporous,  polysporous  and  mono- 
sporous, tho  of  rare  occurrence. 

Spore:  Broad  spindle  shape  with  attenuated  anterior  end.  Shell  rather 
thin.  Each  valve  has  6  to  8  longitudinal  striations  on  the  surface.  A  long 
tail  composed  of  two  halves,  is  developed  at  the  posterior  end.  Two  pyri- 
form  polar  capsules  with  distinctly  visible  coiled  polar  filament  opens  at 
the  anterior  tip.  Sporoplasm,  finely  granular,  contains  an  iodinophilous 
vacuole  which  is  made  distinctly  visible  by  treating  with  Lugol's  solution. 
When  stained  two  typical  nuclei  are  recognized  in  the  sporoplasm.  Dimen- 
sions of  the  fresh  spores:  length  excluding  tail  13.5  to  15/t,  breadth  8  to 
9/x,  thickness  6  to  7.5/i,  length  of  tail  30  to  35/1,  often  up  to  40/i,  polar 
capsule  5  to  6/*  by  3/i,  length  of  polar  filament  40/i. 


Genu 

s     HOFERELLUS     Berg 

1898 

Hoferdlus 

Berg                    1898  :  41 

1898 

Hoferia 

Doflein                1898  :  288-289 

The  characters  of  the  genus  are  described  on  page  59. 
Type  and  only  species:  Hoferellus  cyprini  Doflein. 


1T4  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [412 

HOFERELLUS  CYPRINI  Doflein 
[Figs.  577  to  581] 

1898        Hojeriacy print  Doflein  1898:289-290 

1908        Hoferellus  cyprini  Mercier  1908  :  LIII-LIV 

1910        Hoferellus  cyprini  Plehn  1910  :  20-22 

Habitat:  In  lumen  and  epithelial  cells  of  renal  tubules  of  kidney  of 
Cyprinus  carpio  L.;  France  and  Germany. 

Vegetative  form:  Young  trophozoites  live  in  epithelium.  Adults  free 
in  the  urinary  tubules.  Form  rounded  or  oval.  No  clear  differentiation  of 
protoplasm.  Pseudopodium  unobserved.  Endoplasm  contains  numerous 
granules  and  many  nuclei.  Each  pansporoblast  forms  two  spores.  Smaller 
individuals  20  to  30/*  in  diameter.    Polysporous. 

Spore:  Pyramidal  with  two  short  tail-like  processes  at  the  posterior 
end,  which  are  formed  from  the  shell-valves  like  those  of  Henneguya. 
Between  these  two  processes,  rarely  small  protoplasmic  pointed  processes 
occur.  Each  shell-valve  has  9  to  10  longitudinal  striations  on  it.  Two 
polar  capsules  at  the  anterior  part,  show  clearly  the  coiled  polar  filaments. 
Sporoplasm  has  two  neclei  and  an  io<iinophilous  vacuole.  Dimensions: 
total  length  10  to  \2n,  breadth  8/i,  tail-process  2/i  long,  polar  capsule  3/x  long. 

MYXOSPORIDIA  GENERA  ET  SPECIES  INCERTAE 

Gen.  et  spec,  incert.  Leydig 
1851  Leydig  1851  :  222 

1894        Gen.  et  spec.  incerU  Gurley  1894  :  186 

Habitat:  Cysts  in  the  root  of  tongue  of  Chondrostoma  nasus  L.;  Ger- 
many. 

Gen.  et  spec,  incert.  Leydig 
1851  Leydig  1851  :  222 

1894        Gen.  et  spec,  incert.  Gurley  1894  :  186 

Habitat:  Heart  (auriculo- ventricular  valve)  of  Leuciscus  rutilus  L. 

Gen.  et  spec,  incert.  Leydig 

1851  Leydig  1851  :  223 

1894        Gen.  et  spec,  incert.  Gurley  1894  :  186 

Gen.  et  spec,  incert.  Heckel  et  Kner 
1851  Heckel  and  Kner  1851  :  12 

1894        Gen.  et  spec,  incert.        Gurley  1894  :  186-187 

Habitat:  Branchiae  of  Lucioperca  lucioperca  L.;  Austria. 

Gen.  et  spec,  incert.  Borne 
1886  Borne  1886  :  211 

1894        Gen.  et  spec  incert  Gurley  1894  :  187 

Habitat:  Scomber  scombrus  L. 


4131  STUDIES  ON  MYXOSPORIDIA— KUDO  175 

Genus  incert.  MERLUCII  Perugia 
[Figs.  582  and  583] 
1891        Myxosporidium  merlucii  Perugia  1891  :  22-24 

1894        Myxobolus  ?  merlucii  Gurley  1894  :  242-243 

1899        Myxobolus  merlucii  Labb6  1899  :  100 

Habitat:  Gall-bladder  of  Merlucius  merlucius  L.;  Italy. 

Vegetative  form:  Various  form.  No  differentiation  of  protoplasm. 
Disporous  (?). 

Spore:  Oval,  with  two  polar  capsules. 

Remarks:  The  species  was  placed  in  the  genus  Myxobolus  by  previous 
authors.  The  figures  given  by  Perugia  show  that  the  spores  are  at  least 
dimorphous.  From  the  habitat  and  the  disporous  characters,  one  should 
place  it  rather  in  one  of  the  genera  of  the  Family  Ceratomyxidae. 

Genus  incert.  CONGRI  Perugia 
[Figs.  584  and  585] 
1891        Myxosporidium  congri  Perugia  1891  :  24-25 

1894        Genus  incert.  congri  Gurley  1894  :  182 

1912        Myxobolus  congri  Parisi  1912  :  284 

Habitat:  Gall-bladder  of  Conger  conger  L.;  Genova. 
Vegetative  form :  Floating  in  the  bladder.    Form  variable.    Movements 
incessant,  slow  and  ameboid. 
Spore:  Not  described. 


Gen.  et  spec,  incert.  Linton 

[Fig.  590] 

1891 

Linton 

1891  :  359-361 

1894 

Gen.  et  spec,  incert.                 Gurley 

1894  :  182-183 

Habitat:  Subcutaneous  tissue  of  Notropis  megahps  Raf.;  Ohio  (Black 
River;  September,  October). 

Vegetative  form:  Cysts.  Globular,  discrete  or  aggregated  into  clusters, 
white,  with  minute  patches  of  black  pigment  from  host;  size  varying  from 
2.5mm.  (single  cyst)  to  7mm.  by  5mm.  (clusters) ;  cyst-membrane  composed 
of  connective  tissue. 

Spore:  Top-shaped,  somewhat  flattened;  with  pointed  anterior  and 
broadly  rounded  posterior  end.  Shell  thick,  with  elevated  sutural  ridge. 
Polar  capsules  could  not  be  detected.  Protoplasm  finely  granular.  Dimen- 
sions: length  17/i,  breadth  10/t,  thickness  6;u. 

Remarks :  The  cysts  and  figures  of  spores  given  by  Linton  suggest  that 
it  is  most  probably  a  unicapsular  Myxobolus.  As  Linton  could  not  detect 
(?)  the  polar  capsule,  tho  his  figures  faintly  show  the  said  structure,  it  is 
placed  in  this  group. 


176  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [414 

Gen.  et  spec,  incert.  Mingazzini 
1892  Mingazzini  1892  :  398 

1899  Labbe  1899  :  113 

Habitat:  Ovarian  egg  of  Lacerta  sp. 

Vegetative  form:  Ameboid  with  hyaline  pseudopodia  and  granular 
protoplasm. 

Spore:  Not  observed. 

Gen.  et  spec,  incert.  Nufer 
1905        Myxobolus  sp.  Nufer  1905  :  71,  77,  79,  85, 186 

Habitat:  In  the  connective  tissue  of  branchia  of  Chondrostoma  nasus; 
Lake  of  Lucerne.    A  single  cyst  in  a  single  host  fish. 

Vegetative  form:  Cyst  white,  and  of  1mm.  in  diameter. 

Spore:  With  two  polar  capsules  at  one  pole  and  the  sporoplasm. 
^  Dimensions  or  any  other  characters  are  not  given. 

Remarks:  Altho  Nufer  placed  the  form  in  the  genus  Myxobolus,  this 
must  be  brought  into  the  present  group  in  view  of  the  fact  that  the  iodin- 
ophilous  vacuole  was  not  detected,  and  that  the  observation  is  too  incom- 
plete to  place  it  to  any  one  of  the  genera. 


Gen.  et  spec,  incert.  Mavor 

[Figs.  586  and  587] 

1915 

Mavor 

1915  :  27-28,  32-33 

1916 

Mavor 

1916  :  553-554 

Habitat:  Gall-bladder  of  Urophycis  chuss;  St.  Andrews  (July  to  Sep- 
tember). 

Vegetative  form:  Mavor  writes  as  follows: 

Attached,  usually  in  large  numbers,  to  the  epithelium  of  the  bladder, 
occurs  a  spherical  or  ellipsoidal  trophozoite  which  in  stained  preparations 
is  found  to  contain  numerous  nuclei.  Very  often  clusters  of  Ceratomyxa 
acadiensis  are  found  adhering  to  the  free  surface  of  myxosporidium.  In 
fresh  preparations  the  appearance  is  that  of  budding  from  a  parent  organ- 
ism. An  examination  of  sections  has  shown  a  sharp  division  between  the 
myxosporidium  and  Ceratomyxa  acadiensis. 

Spore:  Not  found. 

Remarks:  Mavor  supposed  that  the  form  under  discussion  probably 
was  some  species  of  Myxidium  or  Chloromyxum. 

Gen.  et  spec,  incert.  Mavor 
[Figs.  588  and  589] 
1916  Mavor  1916a  :  68-69 

Habitat:  Urinary  bladder  of  Stizostedion  vitreum  Mitch.;  Georgian 
Bay  (Canada). 


4151  STUDIES  ON  MYXOSPORJDJA— KUDO  177 

Vegetative  form:  Free  forms  vary  greatly  in  shape,  being  rounded, 
elongated  or  branched.  The  largest  individual  200/*,  Ectoplasm  layer 
clearly  visible,  sometimes  projecting  many  bristle-like  short  processes. 
Endoplasm  contains  greenish  granules.  Trophozoites  also  attached  to 
the  epithelium  by  means  of  deeply  stainable  portion  of  the  body. 

Spore:  Not  observed. 

Remarks:  Mavor  mentions  resemblance  of  the  present  form  to  Myxi- 
dium  lieberkuhni  Biitschli  in  many  respects. 


178  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [416 


KEYS  TO  THE  GENERA  AND  SPECIES  OF  MYXOSPORIDIA 

No  key  to  the  genera  and  species  of  Myxosporidia  has  been  published 
up  to  the  present  time.  This  is  due  of  course  to  the  difficulties  which 
accompany  such  an  attempt.  These  difficulties  lie  chiefly  in  the  incom- 
pleteness of  the  observations  and  descriptions  of  the  majority  of  the  species 
of  Myxosporidia. 

The  writer  has  attempted  in  the  following  pages  to  carry  out  this  task. 
The  key  is  by  no  means  complete,  as  is  unavoidable  in  the  present  state  of 
knowledge  concerning  this  particular  group  of  the  Protozoa. 

Altho  the  spore  is  the  fundamental  factor  used  in  constructing  this  key, 
it  was  necessary  to  refer  also  to  some  other  secondary  characters  such  as 
vegetative  form  and  habitat. 

Some  authors  are  inclined  to  think  that  the  difference  in  host  species 
gives  an  ample  basis  on  which  to  record  the  parasite  as  a  new  species.  In 
some  cases  the  parasite  is  specific  in  a  certain  host  species  while  in  other 
cases  a  number  of  different  host  species  are  infected  by  one  and  the  same 
parasite.  Therefore  one  can  not  lay  much  emphasis  upon  a  difference  of 
hosts  in  fixing  the  identification  of  a  Myxosporidian. 

KEY  TO  THE  GENERA  OF  MYXOSPORIDIA 

1(6)    Spore  approximately  spherical 

Suborder  Sphaerosporea  Kudo  1919 2 

2(3)     Spore  with  four  polar  capsules 

Family  Chloromyxidae  Thdohan  1890 

Genus  Chloromyxum  Mingazzini  1890 (183) 

3(  2)     Spore  with  two  polar  capsules  • 

Family  Sphaerosporidae  Davis  1917 4 

4(  5)     Sutural  line  of  spore  straight 

Genus  Sphaerospora  Thdohan  1892 (185) 

5(  4)     Sutural  line  of  spore  sinuous 

Genus  Sinuolinea  Davis  1917 (186) 

6(  1)    Spore  not  spherical 7 

7(16)    Largest  diameter  of  spore  at  right  angles  to  sutural  line;  two  polar  capsules,  one 
on  each  side  of  sutural  line 

Suborder  Eurysporea  Kudo  1919 

Family  Ceratomyxidae  Doflein  1899 / 8 

8(11)     Shell-valves  prolonged  laterally 9 

9(10)    Shell-valves  hemispherical  or  rounded 

Genus  Leptotheca  Th^lohan  1895 (179) 

10(  9)    Shell-valves  conical;  free  end  tapering  to  a  more  or  less  pointed  end 

Genus  Ceratomyxa  Th61ohan  1892 (180) 

11(8)     Shell- valves  rather  elongated;  circular  in  cross-section 12 

12(13)    Spore  rounded  oblong;  shell  longitudinally  striated;  polar  capsules  pyriform,  with 
or  without  long  and  fine  posterior  filaments 

Genus  Mitraspora  Fujita  1912  emend.  Kudo  1919 (183) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


4171 


STUDIES  ON  MYXOSPORIDIA—KUDO 


179 


13(12 
14(15: 


15(14; 


16(  7 

17(22 

18(21 
19(20; 

20(19; 

21(18; 

22(17 
23(26; 

24(25 

25(24; 

26(23; 

27(30; 
28(29; 

29(28; 
30(27 


Spore  angular,  not  rounded 14 

Spore  pyramidal  in  front  view;  with  its  base  at  anterior  end;  with  or  without  dis- 
tinct anterior  processes;  shell  smooth 

Genus  Myxoproteus  Doflein  1898 (183) 

Spore  isosceles  triangular  in  front  view;  anterior  end  attenuated;  polar  capsules 
spherical  and  large;  shell  with  fine  network-like  ridges;  with  posterior  fringe- 
like processes 

Genus  Wardia  Kudo  1919 (183) 

Largest  diameter  of  spore  coincides  with  or  at  an  acute  angle  to  sutural  plane; 
one  or  two  polar  capsules  which  ^e  in  sutural  plane 

Suborder  Platysporea  Kudo  1919 17 

Spore  fusiform;  two  polar  capsules,  one  at  each  end  of  spore 

Family  MYXiDiiDAETh61ohan  1892 18 

Spore  more  or  less  regularly  fusiform;  shell-valves  symmetrical 19 

Polar  filament  fine  and  long 

Genus  Myxidium  Butschli  1882 (186) 

Polar  filament  thick  and  short 

Genus  Sphaeromyxa  Th^lohan  1892 (188) 

Spore  semi-circular  in  front  view;  polar  filament  fine 

Genus  Zschokkella  Auerbach  1910 (188) 

Spore  not  fusiform;  with  one  or  two  polar  capsules  at  anterior  extremity 23 

Sporoplasm  without  iodinophilous  vacuole 

Family  Myxosomatidae  Poche  1913 24 

Spore  elongated  ovoid  in  front  view;  anterior  end  mostly  pointed 

Genus  Myxosoma  Th61ohan  1892 (189) 

Spore  more  or  less  rounded  in  front  view 

Genus  Lentospora  Plehn  1905 (189) 

Sporoplasm  always  with  iodinophilous  vacuole 

Family  Myxobolidae  Thelohan  1892 27 

Spore  with  posterior  process;  shell  sometimes  striated 28 

Process  more  or  less  long,  projecting  posteriad  along  median  line  of  spore;  process 
either  single  or  double;  shell  sometimes  striated 

Genus  Henneguya  Thelohan  1892 (J^^) 

Process  short  projecting  posteriad  from  sides;  shell  longitudinally  striated 

Genus  Hoferellus  Berg  1898 (173) 

Spore  without  posterior  process;  shell  unstriated;  one  or  two  polar  capsules 

Genus  Myxobolus  Butschli  1882 (189) 


11.    KEY  TO  THE  SPECIES 
•    Genus    LEPTOTHECA    Th61ohan    1895 

1(14)    Spore:  sutural  diameter  always  more  than  half  of  greatest  breadth 2 

2(  7)    Average  sutural  diameter  less  than  10^ 3 

3(  4)    Posterior  margin  of  spore  concave  in  front  view;  sutural  diameter  8  to  9n,  breadth 

12  to  14m.    Trophozoite  usually  with  a  long  process 

Leptotheca  longipes  Auerbach  1910 (63) 

4(  3)    Posterior  margin  of  spore  not  concave 5 

5(  6)     Posterior  margin  of  spore  flattened;  polar  capsules  p3ndform;  sutural  diameter  6 

to  7m,  breadth  11  to  12^.    Trophozoite  with  actively  motile  long  filiform 

pseudopodia  at  rounded  end 

Leptotheca  agilis  Thelohan  1892 (60) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


180  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [418 

6(  5)  Spore  regularly  ovoidal ;  polar  capsules  short  pynf onn,  opening  on  opposite  sides ; 
sutural  diameter  9/u,  breadth  \6y..  Trophozoite  pyriform  without  any  recog- 
nizable pseudopodium 

Leptotheca  fusiformis  Davis  1917 (63) 

7(  2)     Average  sutural  diameter  of  spore  equal  to  or  larger  than  10/* 8 

8(13)     Shell-valves  symmetrically  built;  sutural  ridge  straight 9 

9(10)  Posterior  margin  of  spore  concave  in  front  view;  sutural  diameter  lO/iz,  breadth 
18  to  20/*;  each  spore  formed  independently 

Leptotheca  informis  Auerbach  1910 (63) 

10(  9)     Spore  regularly  ovoidal 11 

11(12)  Trophozoite  extremely  polymorphous.  Spore:  sutural  diameter  10  to  12/«, 
breadth  18  to  20/1 

Leptotheca  polymorpha  Th61ohan  1895 (61) 

12(11)  Typical  form  of  trophozoite  elongated;  anterior  end  depressed  surrounded  by 
short  often  branched  pseudopodia.  Spore:  sutural  diameter  12  to  15/i, 
breadth  18  to  20/t 

Leptotheca  elongata  Thdohan  1895 (60) 

13(  8)  Shell-valves  asymmetrically  built;  sutural  ridge  sinuous;  sutural  diameter  9  to 
lO/i,  breadth  16  to  18/*.    Trophozoite  rounded;  movements  slow 

Leptotheca  lobosa  Davis  1917 (64) 

14(  1)     Sutural  diameter  equal  to  or  less  than  half  of  greatest  breadth 15 

15(18)     Average  sutural  diameter  smaller  than  10/« 16 

16(17)     Spore  arch-shaped  in  front  view;  polar  capsules  pyriform;  sutural  diameter  3  to 
4/*,  breadth  8  to  lO/i 

Leptotheca  parva  Thelohan  1895 (61) 

17(16)     Spore  cylindrical;  sutural  diameter  4.5/t,  breadth  9/» 

Leptotheca  glomerosa  Davis  1917 (65) 

18(15)     Average  sutural  diameter  greater  than  10/* 19 

19(20)  Posterior  margin  of  spore  slightly  concave  in  front  view ;  anterior  end  attenuated ; 
polar  capsules  pyriform;  sutural  diameter  13/*,  breadth  26/t.  Trophozoite 
rounded;  with  active  amoeboid  movements 

Leptotheca  macrospora  Auerbach  1909 (62) 

20(19)  Posterior  margin  of  spore  more  or  less  flattened;  anterior  end  smoothly  rounded; 
polar  capsules  rounded;  sutural  diameter  11/*,  breadth  22 /t.  Trophozoite 
elongated;  pseudopodia  often  anastomose 

Leptotheca  scissura  Davis  1917 (64) 

Incompletely  described  species 

Leptotheca  renicola  Thelohan  1895 (61) 

Leptotheca  hepseti  Thelohan  1895 (62) 

Leptotheca  perlata  Gurley  1894 (62) 

Leptotheca  sp.  Awerinzew  1908 (62) 

Genus    CERATOMYXA    Thelohan     1892 

1(52)     Spore  constant  in  form  and  size 2 

2(21)     Sutural  diameter  equal  to  or  less  than  one-eighth  of  total  breadth 3 

3(10)     Sutural  diameter  not  less  than  one-tenth  of  total  breadth 4 

4(  9)     Pseudopodia  of  vegetative  form  located  at  rounded  end 5 

5(  6)  Pseudopodia  long  filiform;  with  slow  whiplash-like  movements  toward  pointed 
extremity.    Spore:  sutural  diameter  12/t,  breadth  118/* 

Ceratomyxa  flagelUfera  Davis  1917 (77) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named.  ^ 


419J  STUDIES  ON  MYXOSPORIDIA—KUDO  181 

6(  5)     Pseudopodia  short  lobose 7 

7(  8)  Extremities  of  spore  attenuated;  spore  large;  sutural  diameter  10  to  12;*,  breadth 
90  to  100^ 

Ceratomyxa  spkaerulosa  Thdlohan  1892 (66) 

8(  7)  Extremities  of  spore  rounded;  spore  small;  sutural  diameter  4/«,  breadth  34  to 
39m 

Ceratomyxa  slreptospora  Davis  1917 (79) 

9(4)  Pseudopodia  unlocalized ;  from  main  part  of  sporulating  trophozoite  are  branched 
out  from  one  to  six  long  prolongations.  Spore:  sutural  diameter  5  to  7/«, 
breadth  50^ 

Ceratomyxa  a-ppendiculata  Thelohan  1892 (67) 

10  (  3)     Sutural  diameter  equal  to  or  less  than  one- tenth  of  total  breadth 11 

11(14)     Shell-valve  terminating  in  a  fine  thread-like  process  at  distal  end 12 

12(13)     Sutural  diameter  7  to  8/u,  breadth  40  to  50^,  lateral  process  250  to  300/i 

Ceratomyxa  acadiensis  Mavor  1915 (71) 

13(12)     Sutural  diameter  Sfi,  breadth  10  to  12/i,  length  of  lateral  process  20ju 

Ceratomyxa  linospora  Doflein  1898 (69) 

14(11)     Shell-valves  not  terminating  in  thread-like  processes 15 

15(20)     Shell-valve  drawn  out  into  a  delicate  process 16 

16(17)     Lateral  process  ribbon-like;  sutural  diameter  6ft,  breadth  140  to  150m 

Ceratomyxa  taenia  Davis  1917 (74) 

17(16)     Lateral  process  not  ribbon-like,  but  circular  in  cross-section 18 

18(19)  Posterior  margin  of  main  part  of  spore  flattened;  sutural  diameter  12m,  breadth 
115  to  140m;  trophozoite  disporous 

Ceratomyxa  sphairophora  Davis  1917 (73) 

19(18)  Posterior  margin  of  main  part  of  spore  rounded;  sutural  diameter  7m,  breadth 
80m.    Trophozoite  monosporous  or  disporous 

Ceratomyxa  spinosa  Davis  1917 (80) 

20(15)  Shell-valve  tapering  gradually  to  attenuated  point;  asjoimietrical;  sutural  diame- 
ter 9m,  breadth  115m 

Ceratomyxa  attenuata  Davis  1917 (75) 

21(  2)     Sutural  diameter  more  than  one-eighth  of  total  breadth 22 

22(45)     Sutural  diameter  equal  to  or  more  than  one-fifth  of  breadth 23 

23(34)     Shell-valves  symmetrically  built 24 

24(29)     Shell-valves  attenuated  at  distal  end 25 

25(26)  Pseudopodia  peculiar  network-like  foim.  Spore:  sutural  diameter  12  to  20m, 
breadth  50  to  80m 

Ceratomyxa  ramosa  Awerinzew  1907 (69) 

26(25)     Pseudopodia  never  unite  together 27 

27(28)  Shell-valves  curved  greatly  posteriad;  polar  capsules  rounded;  sutural  diameter 
8  to  9m,  breadth  16(?)m.    Trophozoite  elongated  pyriform. 

Ceratomyxa  recurvata  Davis  1917 (75) 

28(27)  Shell-valves  not  curved;  two  thickenings  on  posterior  margin  equidistant  from 
sutural  line;  polar  capsules  pyrifonn;  sutural  diameter  40  to  45m,  thickness  25 
to  30m,  breadth  124  to  140m 

Ceratomyxa  tylosuri  Awerinzew  1913 (70) 

29(24)     Shell-valve  rounded  at  distal  end 30 

30(31)     Spore  arch-shaped;  sutural  diameter  and  thickness  12  to  15m,  breadth  50  to  60m. 
Trophozoite  large  amoeboid. 

Ceratomyxa  spari  Awerinzew  1913 (70) 

31(30)     Spore  straight 32 

Numbers  eiiclose(}  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


182  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [420 

32(33)    Shell-valves  shorter  (sutural  diameter:  breadth  =  1:1.6) 

Ceratomyxa  coris  Georg6vitch  1916 (72) 

33  (32)     Shell-valves  longer  (sutural  diameter :  breadth = 1 :2 . 6) 

Ceratomyxa  herouardi  Georgevitch  1916 (72) 

34(23)     Shell-valves  as3Tmnetrically  built 35 

35(40)     Spore  arch-shaped  in  front  view 36 

36(37)  Sutural  diameter  equal  to  one-fifth  of  breadth;  sutural  diameter  lO/t,  breadth 
50m 

Ceratomyxa  globulifera  Thelohan  1895 (67) 

37(36)     Sutural  diameter  more  than  one-fifth  of  total  breadth 38 

38(39)     Spore:  breadth  shorter;  sutural  diameter  14m,  breadth  17m 

Ceratomyxa  abbreviata  Davis  1917 (76) 

39(38)     Spore:  breadth  longer;  sutural  diameter  12  to  15m,  breadth  45  to  50m 

Ceratomyxa  reticularis  Thelohan  1895 (68) 

40(35)     Spore  straight 41 

41(42)    Sutural  diameter  11m,  breadth  27m.    Trophozoite  alwajrs  roimded 

Ceratomyxa  amorpha  Davis  1917 (78) 

42(41)     Sutural  diameter  6m 43 

43(44)    Trophozoite  with  active  pseudopodia.     Spore:  sutural  diameter  6m,  breadth 

22  to  24m 

Ceratomyxa  undtdala  Davis  1917 (79) 

44(43)  Trophozoite  with  inactive  pseudopodia.  Spore:  sutural  diameter  6m,  breadth 
31m 

Ceratomyxa  inaequalis  Doflein  1898 (68) 

45(22)     Sutural  diameter  less  than  one-fifth  of  total  breadth 46 

46(49)    Breadth  of  Sf)ore  equal  to  or  greater  than  50m 47 

47(48)  Shell-valve  tapering  gradually  toward  distal  end;  sutural  diameter  8m,  breadth 
50  to  56m.    Trophozoite  usually  elongated  pyriform 

Ceratomyxa  mesospora  Davis  1917 (73) 

48(47)  Shell-valve  rounded  at  distal  end;  sutural  diameter  6  to  7m,  breadth  50m.  Tro- 
phozoite usually  rounded  or  irregular  form;  size  small 

Ceratomyxa  aggregate  Davis  1917 (79) 

49(46)     Breadth  of  spore  smaller  than  30m SO 

50(51)  Trophozoite  ordinarily  spherical,  diameter  not  exceeding  16  to  20m;  protoplasm 
extremely  pale  looking.    Spore :  sutural  diameter  5m,  breadth  25  to  30m. 

Ceratomyxa  pallida  Thelohan  1895 (67) 

51(50)  Trophozoite  pyriform  with  a  long  posterior  process;  movements  by  wavelike 
motion  of  ectoplasm;  also  active  backward  movements  of  pseudopodia.  Spore 
asymmetrically  built;  sutural  diameter  5m,  breadth  24  to  28m 

Ceratomyxa  agglomerata  Davis  1917 (77) 

52(  1)     Spore  variable  in  size  and  form 53 

53(54)  Variation  in  number  of  shell-valves  conspicuous;  sutural  diameter  5m,  breadth 
25m 

Ceratomyxa  truncata  Thelohan  1895 (67) 

54(53)    Variable  in  size  and  form  of  spore,  but  not  in  niunber  of  shell- valve 55 

55(60)    Trophozoite  more  or  less  definite  in  shape 56 

56(59)    Trophozoite  usually  pyriform 57 

57(58)    Trophozoite  disporous.    Spore:  sutural  diameter  7  to  9m,  breadth  15  to  38m 

Ceratomyxa  lunata  Davis  1917 (76) 

58(57)  Trophozoite  monosporous  or  disporous.  Spore:  sutural  diameter  5  to  6m, 
breadth  18  to  25m 

Ceratomyxa  monospora  Davis  1917 (78) 

Niunbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


4211  STUDIES  ON  MYXOSPORIDIA—KUDO  183 

59(56)  Trophozoite  always  rounded,  never  pyrifonn.  Spore:  sutural  diameter  6/«, 
breadth  \(>n 

Ceratomyxa  navicularia  Davis  1917 (80) 

60(55)    Trophozoite  polymorphous 61 

61(62)     Shell-valves  symmetrically  built;  sutural  diameter  5  to  8^,  breadth  20  to  31/a 

Ceratomyxa  arcuata  Thelohan  1892 (65) 

62(61)  Shell- valves  often  asymmetrically  built;  sutural  diameter  8  to  14^,  breadth  SO  to 
80/i 

Ceratomyxa  drepanopsettae  Awerinzew  1907 (70) 

Incompletely  described  species 

Ceratomyxa  sp.  (?)  Awerinzew  1913 (71) 

Ceratomyoca  sp.  (?)  Awerinzew  1913 (71) 

Ceratomyxa  sp.  Georg6vitch  1916 (72) 

Genus    MYXOPROTEUS    Doflein    1898 
1(  2)     Spore  with  two  long  (5ju)  processes  extending  anteriad  from  sides;  sutural  diameter 
9 IX,  breadth  12  m 

Myxoproteus  cornuttis  Davis  1917 (82) 

2(  1)     Spore  without  long  process 3 

3(  4)  Spore  with  two  small  spinous  processes  at  anterior  end;  sutural  diameter 
25  m,  breadth  18  to  20/* 

Myxoproteus  ambiguus  Thelohan  1895 (81) 

4(  3)  Spore  without  any  process;  posterior  end  slightly  pointed;  sutural  diameter  12^, 
breadth  10  to  11m 

Myxoproteus  cordiformis  Davis  1917 (81) 

Genus    WARDIA    Kudo    1919 
1  Spore  isosceles  triangular  form;  shell  with  network-like  striations  which  end  in 

fringe-like  processes  at  posterior  margin;  sutural  diameter  9  to  10m,  breadth  10 
to  12m,  diameter  of  polar  capsule  4m. 

Wardia  ovinocua  Kudo  1919 (82) 

Doubtfully  placed  species 

Wardia  ohlmacheri  (Gurley  1894) (83) 

Genus    MITRASPORA    Fujita    1912  emend.    Kudo    1919 

1(  4)     Spore  with  posterior  filaments 2 

2(  3)  Posterior  filaments  short  (5  to  6m  long);  length  10m,  breadth  8  to  9m,  thickness 
6  to  8m,  polar  capsule  4m  by  1.5  to  2m 

Mitraspora  cyprini  Fujita  1912 ! (84) 

3(  2)  Poisterior  filaments  of  spore  long  (up  to  28m)  ;  length  10  to  11m>  polar  capsule  4  to 
4.5m  long 

Mitraspora  caudata  Parisi  1910 (85) 

4(  1)  Spore  without  posterior  filament;  anterior  end  slightly  attenuated;  posterior  end 
truncate;  length  15  to  17m,  breadth  5  to  6m,  thickness  4.5  to  5.5m,  polar  cap- 
sule 7.5m  by  2m 

Mitraspora  elongala  Kudo  1919 (85) 

Genus    CHLOROMYXUM    Mingazzini    1890 

1(  4)     Spore  with  posterior  appendage 2 

2(  3)  Posterior  appendage  fine  and  numerous;  length  6  to  9ii.,  breadth  5  to  6m,  polar 
capsule  2  to  3  m  by  1  to  2  m 

Chloromyxum  leydigiMrngSiZzmi  1890 * (87) 

Numbers  enclosed  in  parentheses  refei  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


184  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [422 

3(  2)    Posterior  appendage  single  or  bifurcated;  length  8^,  breadth  6  to  7^,  appendage 
10/t  long 

Chloromyxum  caudatum  Th61ohan  1895 (88) 

4(  1)     Sp)ore  without  posterior  appendage 5 

5(34)    Spore  circular  or  subcircular  in  front  view;  parasitic  in  body  cavity  of  host 6 

6(29)     Shell-valves  marked  with  striations  or  ridges 7 

7(24)    Main  part  of  striations  or  ridges  parallel  to  sutural  line 8 

8(11)     Shell-valves  partially  marked 9 

9(10)     Ridges  on  each  shell- valve  variable  in  number  (six  found  in  original  drawing) 
ruiming  closely  to  sutural  line;  diameter  lO.Sfi 

Chloromyxum  duhium  Auerbach  1908 (91) 

10(  9)  Each  shell-valve  with  one  ridge  from  which  eight  to  twelve  short  ones  are  directed 
toward  centre  of  valve;  oval  in  profile;  length  and  breadth  8  to  lO/z,  thickness 
5  to  7m 

Chloromyxum  trijugum  Kudo  1919 (96) 

11(  8)     Entire  shell-valve  marked 12 

12(19)     Shell-valve  marked  with  fine  striations 13 

13(16)     Spore  oval  in  lateral  view 14 

14(15)  Trophozoite  larger;  size  up  to  SOn  by  20/*;  polysporous  (up  to  eight  spores)  rarely 
disporous.    Spore:  length  8  to  9/x,  breadth  6  to  7/x,  thickness  5  to  6ft 

Chloromyxum  misgurni  Kudo  1916 (93) 

15(14)  Trophozoite  smaller;  size  up  to  35m  in  diameter;  polysporous  (up  to  six  spores) 
or  disporous.    Spore:  length  8m,  breadth  7m,  thickness  5  to  6m 

Chloromyxum  catostomi  Kudo  1919 (98) 

16(13)     Spore  circular  in  lateral  view 17 

17(18)  Trophozoite  rounded;  40  to  45m  by  28  to  40m.  Spore:  diameter  10  to  13m,  polar 
capsule  4  to  6m  long 

Chloromyxum  protei  Joseph  1905 (90) 

18(17)  Trophozoite  irregular  form;  33  to  35m  in  average  length.  Spore:  striations 
thicker  and  somewhat  wavy;  diameter  9  to  9 .  5m,  polar  capsule  3n  long 

Chloromyxum  thymalli  Lebzelter  1912 (9Z) 

19(12)     Shell-valves  marked  with  ridges 20 

20(21)  Trophozoite  small  (average  diameter  of  adults  about  20m)  5  monosporous,  rarely 
disporous.  Spore:  shell- valves  with  ridges  marked  antero-posteriad;  diameter 
10  to  11m 

Chloromyxum  crislatum  Leger  1906 (91) 

21(20)     Trophozoite  large,  diameter  reaching  40  to  50m 22 

22(23)  Ridges  on  shell-valves  united  into  a  line  at  each  end  and  unequal  in  thickness; 
spore  small;  length  10  to  12m,  breadth  8  to  10m 

Chloromyxum  fujitai  Kudo  1916 (93) 

23(22)  Shell-valve  with  two  circular  and  two  small  ridges;  spore  large;  length  16m, 
breadth  10m 

Chloromyxum  koi  Fujita  1913 (92) 

24(  7)     Striations  or  ridges  not  parallel  to  sutural  line 25 

25(26)     Striations  irregular;  posterior  margin  thickened  at  sides;  diameter  7 . 5  to  9m 

Chloromyxum  wardi  Kudo  1919 (99) 

26(25)     Striations  parallel  to  each  other < 27 

27(28)     Striations  forming  acute  angles  with  sutural  line;  diameter  8  to  9m 

Chloromyxum  truttae  L^ger  1906 (90) 

28(27)  Four  ridges  on  posterior  half  of  shell- valve  converging  toward  anterior  end; 
diameter  7  m 

Chloromyxum  granulosum  Davis  1917 (96) 

Niunber  senclosed  in  parentheses  refer  to  the  page  of  the  article  aa  which  is  found  the  description  of  the  spedes 
named. 


423]  STUDJES  ON  MYXOSPORIDIA—KUDO  185 

29(  6)    Shell-valves  without  marking,  beside  sutural  ridge 30 

30(31)  Anterior  end  of  spore  rounded;  diameter  7  to  S/z;  one  or  two  short  spinous  thick- 
enings at  posterior  margin 

Chloromyxum  fluviatile  Thdlohan  1892 (89) 

31(30)    Anterior  end  of  spore  mucronate  or  truncate 32 

32(33)     Anterior  end  of  spore  mucronate;  length  8m 

CMorotnyxum  mucronatum  Gurley  1893 (89) 

33(32)     Anterior  end  of  spore  truncate;  spore  large;  length  40  to  48 /x,  breadth  30  to  38^ 

Chloromyxum  magnum  Awerinzew  1913 (92) 

34 (  5)  Spore  rounded  quadrangular  in  end  view;  conical  in  front  view;  parasitic  in  mus- 
cular tissue  of  fish 35 

35(36)     Length  of  spore  larger  than  breadth;  length  6/i,  breadth  5m 

Chloromyxum  quadratum  Thelohan  1895 (88) 

36(35)     Length  (sutural  diameter)  of  spore  smaller  than  breadth 37 

37(38)  Spore  variable  in  form;  anterior  end  narrower  or  broader  than  posterior  end; 
length  4  to  4.75m,  breadth  5.4  to  6.5m 

Chloromyxum  clupeidae  Hahn  1917 (94) 

38(37)  Anterior  end  of  spore  drawn  out;  almost  circular  in  end  view;  length  6m,  breadth 
7.5m 

Chloromyxum  funduli  Hahn  1915 (93) 

Incompletely  described  species 

Chloromyxum  diploxyi  Gurley  1893 (90) 

Chloromyxum  sp.  Awerinzew  1908 (91) 

Genus  SPHAEROSPORA    Thflohan     1892 

1(8)     Shell-valve  of  spore  without  marking  except  sutural  ridge 2 

2(  7)    Vegetative  form  amoeboid 3 

3(  4)  Movements  of  vegetative  form  active.  Spore:  sutural  ridge  fairly  well  marked; 
a  pair  of  short  filaments  become  visible  at  anterior  end  on  warming;  di- 
ameter 8m 

Sphaerospora  masovica  Cohn  1902 (101) 

4(  3)     Vegetative  form  without  active  movements 5 

5(  6)  Spore:  sutural  ridge  not  prominent;  polar  capsule  pyriform;  diameter  8  to  13Mf 
polar  capsule  4  to  5m  by  2 . 5  to  3 .  5m 

Sphaerospora  carassii  Kudo  1919 (103) 

6(  5)  Spore:  sutural  ridge  prominent;  polar  capsule  spherical;  slightly  attenuated  at 
anterior  end;  diameter  10  to  11m 

Sphaerospora  elegans  Thelohan  1892 (100) 

7(  2)     Vegetative  form  produces  cyst  in  tissue.    Spore:  diameter  8  to  9m 

Sphaerospora  platessae  Woodcock  1904 (102) 

8(  1)     Shell-valves  striated 9 

9(10)    Polar  capsules  divergent;  diameter  of  spore  10  to  12m,  thickness  8m 

Sphaerospora  diver  gens  Thdlohan  1895 . .  ; (100) 

10(  9)     Polar  capsules  not  divergent 11 

11(14)     Striation  marked  antero-posteriad 12 

12(13)  Spore  with  a  quadrangular  lamella  at  anterior  margin;  striations  ending  in  small 
spines  at  posterior  margin;  length  12  to  14m,  breadth  10  to  12m 

Sphaerospora  rostrata  Th61ohan  1895 (101) 

13(12)    Spore  smooth-contoured;  polar  capsules  parallel  to  each  other;  diameter  7  to  10m 

Sphaerospora  polymorpha  Davis  1917 (102) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


186  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [424 

14(11)    Faint  concentric  striations;  pointed  at  sides  and  middle  part  of  posterior  margin; 
polar  capsules  unequal  in  size;  length  7  to  8m,  breadth  6  to  7ai,  thickness  Sn 

Splioerospora  angulata  Fujita  1912 (102) 

Incompletely  described  species 

Sphaerospora  sp.  Davis  1917 (103) 

Sphaerospora  sp.  Southwell  et  Prashad  1918 (103) 

Genus    SINUOLINEA    Davis    1917 

1(  4)    Spore  with  two  processes 2 

2(  3)  Processes  lateral  and  long  (20m);  spore:  9  to  lift  long,  9^  broad,  process  18  to 
22ft  long 

Sinnolinea  brachiophora  Davis  1917 (106) 

3(  2)  Processes  posteriad  from  sides  and  short;  diameter  12  to  13^.  Trophozoite 
opaque 

Sinuolinea  opacita  Davis  1917 (106) 

4(  1)    Spore  without  process 5 

S(  6)  Trophozoite  with  active  amoeboid  movements.  Spore:  sutiural  ridge  S-shaped 
at  anterior  part;  length  ISm,  breadth  12m,  thickness  8m 

Sintiolinea  arborescens  Davis  1917 (105) 

6(  5)    Trophozoite  with  slow  amoeboid  movements ■ 7 

7(  8)  Sutural  plane  much  twisted  on  its  axis;  capsulogenous  cells  large  occupying  more 
than  half  of  sporal  cavity;  polar  capsules  opening  on  opposite  sides;  diameter 
12  to  14m 

Sinuolinea  capsularis  Davis  1917 (lOS) 

8(  7)    Sutural  plane  not  much  twisted;  diameter  ISm 

Sinuolinea  ditnorpha  Davis  1916 (104) 

Genus    MYXIDIIIM    ButschU    1882 

1(16)    Breadth  of  spore  equal  to  or  more  than  half  of  length 2 

2(  7)     Shell-valves  imstriated 3 

3(  6)     Sutural  plane  curved  into  anS 4 

4(  5)    Spore  small;  length  8  to  12m,  breadth  4  to  6m 

Myxidium  incurvatum  Thelohan  1892 (108) 

5(  4)     Spore  large;  much  broader;  length  20.8  to  23.4m,  breadth  13  to  15.6m 

Myxidium  inflatum  Auerbach  1909 (HI) 

6(  3)    Sutural  plane  straight;  spore  cylindrical;  surroimded  by  a  gelatinous  envelope; 

length  10  to  11m,  breadth  6m 

Myxidium  glutinosum  Davis  1917 (115) 

7(  2)    Shell-valves  striated 8 

8(  9)    Sutural  line  curved  into  an  S;  form  oval;  circular  in  cross-section;  openings  of 

polar  capsules  pointed;  length  11  to  13m,  breadth  8  to  9m 

Myxidium  oviforme  Parisi  1912 (114) 

9(  8)    Sutural  line  straight 10 

10(13)    Sutural  line  coincides  with  longitudinal  axis  of  spore 11 

11(12)    Sutural  ridge  distinct;  extremities  mucronate;  length  9  to  10m,  breadth  5  to  5 .  6m, 

thickness  4 .  75  to  5  m.   Vegetative  form  produces  cysts,  800  to  900m  in  diameter 

Myxidium  giardi  Cep^de  1906 (HO) 

12(11)    Sutural  ridge  faintly  marked;  extremities  gradually  drawn  out;  length  11m, 

breadth  8m.    Trophozoite  large  and  leaf-like;  diameter  up  to  1 .35  mm. 

Myxidium  phyllium  Davis  1917 (116) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


425]  STUDIES  ON  MYXOSPORIDIA— KUDO  '         187 

13(10)     Sutural  line  forming  an  acute  angle  with  longitudinal  axis  of  spore 14 

14(15)     Shell  thickened  at  extremities;  polar  capsules  ovdidal;  length  10  to  14m,  breadth 
6  to  8iu,  length  of  polar  capsule  4^ 

Myxidium  striatum  Cunha  et  Fonseca  1917 (116) 

15(14)     Shell  uniformly  thick;  polar  capsules  rounded  pyriform;  length  10  to  12^,  breadth 
6n,  length  of  polar  capsule  3  to  4/i 

Myxidium  macrocapsulare  Auerbach  1910 (113) 

16(  1)    Breadth  of  spore  less  than  half  of  length 17 

17(34)    Breadth  more  than  one-third  of  length 18 

18(25)     Shell-valves  unstriated 19 

19(22)    Extremities  of  spore  ported 20 

20(21)     Spore:  extremities  sharply  pointed;  greatly  curved;  narrow;  length  12  to  14/x, 
breadth  5 . 5  to  6ft,  thickness  2.5  to  3  ft 

Myxidium  depressum  Parisi  1912 (114) 

21(20)     Spore:  extremities  not  so  sharply  pointed;  not  greatly  curved;  broader;  length 
16.2  to  19m,  breadth  7  to  9m 

Myxidium  bergense  Auerbach  1909 (112) 

22(19)    Extremities  of  spore  not  pointed 23 

23(24)     Spore  larger;  length  15  to  20m,  breadth  7  to  8m 

Myxidium  sphaericum  Th^lohan  1895 (109) 

24(23)     Spore  smaller;  length  6(?)  to  14m,  breadth  4  to  6ft.   Trophozoite  mictosporous 

Myxidium  gadi  Georg^vitch  1916 (115) 

25(18)     Shell-valves  striated 26 

26(33)     Spore  definite  in  shape 27 

27(28)     Spore  constricted  in  middle  part  of  length;  length  15m 

Myxidium  histophilum  Th^lojian  1895 (109) 

28(27)     Spore  regularly  fusiform 29 

29(30)    Vegetative  form  produces  cyst.    Spore:  length  12  to  15m,  breadth  6m 

Myxidium  harhatulae  C6pede  1906 (HO) 

30(29)     Vegetative  form  does  not  produce  cyst 31 

31(32)     Sutural  line  slightly  curved  in  S-form;  length  15  to  16m,  breadth  and  thickness 

5.5  to  6m 

Myxidium  americanum  Kudo  1919 (117) 

32(31)    Sutural  line  not  curved  in  S-shape,  but  bent  to  one  side;  length  15  to  18m,  breadth 
and  thickness  6  to  7  m 

Myxidium  kagayamai  Kudo  1919 (117) 

33(26)     Spore  variable  in  form;  straight  and  constricted;  one  side  concave,  the  other  con- 
vex; arch-shaped,  etc.;  length  13  to  18m,  breadth  5.2  to  5.8m 

Myxidium  pfeifferi  Auerbach  1908 (HI) 

34(17)     Breadth  of  spore  equal  to  or  less  than  one-third  of  length 35 

35(40)     Shell-valves  unstriated 36 

36(37)     Spore  greatly  elongated  (breadth:  length  =  1:6. 2);  length  2 1.6  to  25.  2  m,  breadth 

3.6  to  4m 

Myxidium  procerum  Auerbach  1910 (112) 

37(36)    Spore  less  elongated  (breadth:  length  =  l:3  or  1:3.4) 38 

38(39)     Spore  large;  valves  asymmetrical;  length  28m,  breadth  8m 

Myxidium  giganleum  DoiBein  1898 (IK^) 

39(38)     Spore  small;  valves  symmetrical;  length  12m,  breadth  3  to  4m 

Myxidium  danilewskyi  Laveran  1897 (109) 

40(35)     Shell-valves  striated 41 

41(44)     Spore  definite  in  shape 42 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


188  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [426 

42(43)    Length  of  polar  capsule  more  than  one-fourth  of  that  of  spore;  spore  18  to  20^ 
long,  5  to  6m  broad 

Myxidium  lieberkuhni  Biitschli  1882 (107) 

43(42)    Length  of  polar  capsule  less  than  one-seventh  of  that  of  spore;  length  16  to  17/x, 
breadth  5^ 

Myxidium  mackiei  Bosanquet  1910 (112) 

44(41)     Spore  variable  in  shape;  S-form,  straight  fusiform,  etc.;  length  9.1/u,  breadth 
2 .  8fx.    Vegetative  form  produces  cyst 

Myxidium  anguillae  Ishii  1915 (1^4) 

Incompletely  described  species 

Myxidium  sp.  Gurley  1894 (109) 

Myxidium  sp.  Awerinzew  1908 (H^) 

Myxidium  sp.  Mavor  1915 (115) 

Genus    SPHAEROMYXA    Th61ohan     1892 

1(  6)     Spore  straight,  not  arch-shaped 2 

2(  5)     Shell-valves  symmetrical 3 

3(4)     Ends  of  spore  truncate ;  stria tions  longitudinal ;  length  1 5  to  20//,  breadth  5  to  6/* . 

Sphaeromyxa  balbianii  Thelohan  1892 (H^) 

4(  3)     Ends  of  spore  rounded;  sutural  plane  forming  some  angles  with  longitudinal  axis 
of  spore;  striations  transverse;  length  12  to  14/x,  breadth  9  to  10m 

Sphaeromyxa  immersa  Lutz  1889 (119) 

5(  2)     Shell-valves  asymmetrical;  unstriated;  ends  less  truncate;  dimensions  about 
twice  or  three  times  larger  than  those  of  Sphaeromyxa  balbianii 

Sphaeromyxa  gaskrostei  Georg^vitch (121) 

6(  1)     Spore  arch-shaped,  not  straight. .  ^ 7 

7(  8)     Shell-valves  indistinctly  striated;  ends  truncate;  length  22  to  28m,  breadth  3  to 
4.3m 

Sphaeromyxa  sabrazesi  Laveran  et  Mesnil  1900 (120) 

8(  7)     Shell-valves  unstriated 9 

9(10)     Spore  extremely  large;  length  75  to  80m,  breadth  18  to  20m;  ends  slightly  tapering 

Sphaeromyxa  exneri  Awerinzew  1913 (121) 

10(  9)     Spore  less  than  35m  in  length 11 

11(12)     Extremities  rounded;  length  30  to  35m,  breadth  8m 

Sphaeromyxa  incurvata  Doflein  1898 (119) 

12(11)     Extremities  truncate;  sutural  ridge  often  twisted  in  S-form;  length  20.8  to  26m, 
breadth  and  thickness  5 .  4m 

Sphaeromyxa  hellattdi  Auerbach  1909 (121) 

Genus    ZSCHOKKELLA    Auerbach     1910 

1(  4)     Shell-valves  unstriated 2 

2(  3)    Openings  of  polar  capsules  on  flattened  side;  spore  large;  length  16  to  28.8m, 
breadth  13  to  18m 

Zschokkella  hildae  Auerbach  1910 (122) 

3(  2)    Openings  o^  polar  capsules  at  pointed  ends;  spore  small;  length  11m,  breadth  7m 

Zschokkella  globulosa  Davis  1917 (123) 

4(  1)     Shell-valves  striated 5 

5(  6)    Openings  of  polar  capsules  at  pointed  ends;  polar  capsules  spherical;  spore  larger; 
length  10  to  14m,  breadth  6  to  7m 

Zschokkella  acheUognathi  Kudo  1916 (123) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  b  found  the  description  of  the  species 
named. 


427]  STUDIES  ON  MYXOSPORIDIA— KUDO  189 

6(  5)  Openings  of  polar  capsules  on  side;  polar  capsules  rounded  pyriform;  spore 
smaller;  length  9.5  to  11. 5^,  breadth  6.5  to  7^ 

Zschokkella  nova  Klokacewa  1914 (122) 

Genus    MYXOSOMA    Th^lohan    1892 
1(  2)     Spore:  shell  thickened  at  anterior  end;  length  12  to  \d>ix,  breadth  7  to  8/*,  polar 
capsule  6  to  7/t  by  2  to  3/i.    Cysts  polymorphous 

Myxosoma  dujardini  Thelohan  1892 (124) 

2(  1)  Spore:  shell  of  uniform  thickness  and  with  seven  to  ten  folds  on  suturaledge; 
length  14^*,  breadth  8m,  thickness  6^,  polar  capsule  8/*  by  2>i.  Cysts  spherical 
up  to  360 jLi  in  largest  diameter 

Myxosoma  fundtdi  Kudo  1918 (125) 

Ambiguous  form 

Myxosoma  lobatutn  Nemeczek  1911 (124) 


1(8: 

2(  3 


3(  2 
4(5: 


5(4 
6(7 


7(6: 
8(1 

9(io: 

10(  9 


1(18 
2(  9 
3(  6 

4(5 


5(4: 


6(  3 

7(  8: 


Genus    LENTOSPORA    Plehn     1905 

Spore  circular  in  front  view 2 

Vegetative  form  produces  cysts.    Spore:  length  and  breadth  6.3  to  Tn,  thick- 
ness 4.2  to4.9/t 

Lentospora  dermatobia  Ishii  1916 (1^) 

Vegetative  form  does  not  produce  cysts  or  cysts  unobserved 4 

Spore  small;  trophozoites  found  in  the  blood  vessel  of  the  brain.    Spore  5  to  5.5/« 
in  diameter. 

Lentospora  encephalina  Mulsow  1911 (126) 

Spore  large,  greater  than  7 .  S^i  in  average  diameter 6 

Spore  slightly  pointed  at  anterior  end;  length  8  to  10/t,  breadth  7  to  8iu,  thickness 
5  to6/n 

Lentospora  acuta  Fujita  1912 (127) 

Anterior  end  of  spore  roimded;  diameter  6  to  lO/i 

Lentospora  cerebralis  Plehn  1905 (125) 

Spore  oval  in  front  view 9 

Spore  symmetrically  built;  length  12/a,  breadth  9.5/i,  thickness  6>x 

Lentospora  multiplicata  Reuss  1906 (126) 

Spore  asymmetrically  built;  length  10  to  ll/x,  breadth  6.5  to  7/z 

Lentospora  asymmetrica  Parisi  1912 (126) 


Genus    MYXOBOLUS    Butschli     1882 

Spore  with  one  polar  capsule 2 

Breadth  of  spore  equal  to  or  more  than  half  of  length ; 3 

Breadth  of  spore  equal  to  half  of  length 4 

Spore  larger;  often  calabash-shaped;  anterior  end  drawn  out  into  a  rounded  tip; 
shell  thickened  at  tip;  length  15^i,  breadth  7  to  8/z,  thickness  5  to  6/i 

Myxobolus  toyamai  Kudo  1915 (131) 

Spore  smaller;  anterior  end  pointed;  shell  of  uniform  thickness;  length  9  to  10|«, 
breadth  4.5  to  5.5|i,  thickness  Six 

Myxobolus  oculi-leucisci  Trojan  1909 (130) 

Breadth  of  spore  more  than  two-thirds  of  length 7 

Polar  capsule  small  and  oblique  in  position 

Myxobolus  unicapsulatus  Gurley  1893 (12i^) 

8(  7)     Polar  capsule  long  and  median  in  position;  spore  broader;  length  13 . 2  to  13 . 6/t, 
breadth  10.1  to  10.3^1 

Myxobolus  sent  Southwell  et  Prashad  1918 (132) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


190  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [428 

9(  2)    Breadth  of  spore  less  than  half  of  length 10 

10(17)    Breadth  of  spore  more  than  two-fifths  of  length 11 

11(16)     Spore  without  any  process 12 

12(15)     Shell  of  uniform  thickness 13 

13(14)  Spore  bent  to  one  side;  shell  thickened  at  slightly  rounded  anterior  tip;  sutural 
edge  without  marking;  length  16  to  18/z,  breadth  7  to  8/tt,  polar  capsule  5 
to  7  m  long 

Myxobolm  piriformis  Th61ohan  1892 (129) 

14(13)  Spore  straight;  shell  thickened  at  posterior  part;  sutural  edge  with  four  to  six 
markings;  length  18  to  20)Lt,  breadth  8^,  thickness  6)u,  polar  capsule  9  to  10^ 
long 

Myxobolus  fuhrmanni  Auerbach  1909 (130) 

15(12)  Shell  of  uniform  thickness;  valves  symmetrical;  sutural  edge  with  markings  up  to 
12  in  number;  spore  14  to  15 .  5ju  long,  6  to  7 . 3/x  broad,  5  to  6m  thick,  polar  cap- 
sule 6.3m  by  2  to  3m 

Myxobolus  misgurni  Kudo  1919 (133) 

16(11)  Spore  with  a  posterior  process,  5m  in  length  and  as  broad  as  spore;  length  17  to 
18m,  breadth  7 .5  to  8m,  polar  capsule  7m  by  4m 

Myxobolus  notatus  Mavor  1916 (131) 

17(10)  Breadth  of  spore  about  one-fourth  of  length;  spore  large;  polar  capsule  extremely 
large;  length  30  to  32m,  breadth  7  to  8m,  length  of  polar  capsule  22  to  23m 

Myx<jibolus  rohitae  Southwell  et  Prashad  1918 (132) 

18(  1)     Spore  with  two  polar  capsules 19 

19(24)     Form  of  spore  variable 20 

20(23)     Spore  with  an  intercapsular  appendix  at  anterior  end 21 

21(22)  Spore  oval;  length  10  to  12m,  breadth  8  to  9m,  thickness  6m,  polar  capsule  5m  by  2 
to  3m 

Myxobolus  midleri  BUtschli  1882 (128) 

22(21)  Spore  pyriform  or  elongated  oval;  length  11  to  16m,  breadth  8  to  13m,  polar  cap- 
sule 6m  by  4m 

Myxobolus  cydoides  Gurley  1893 (140) 

23(20)  Spore  without  intercapsular  appendix;  circular  form  7  to  8m,  breadth  8  to  10m, 
thickness  6m,  polar  capsule  4  to  5m  by  2m 

Myxo6o/Mj  Ay/a€  Johnston  et  Bancroft  1918 (1S3) 

24(19)    Form  of  spore  definite 25 

25(28)     Polar  capsules  in  each  spore  regularly  of  considerably  different  size 26 

26(27)  Spore  with  an  intercapsular  appendix;  anterior  end  roimded;  sutural  edge  with 
folds  (3  to  5);  length  10  to  12m,  breadth  8m 

Myxobolus  dispar  Th^lohan  1895 (135) 

27(26)  Spore  without  intercapsular  appendix;  anterior  end  pointed;  no  fold  on  sutural 
edge 

Myxobolus  inaequalis  Gurley  1893 (135) 

28(25)     Polar  capsules  approximately  of  equal  form  and  size 29 

29(30)     Sutural  diameter  smaller  than  breadth;  length  6  to  7m,  breadth  8/i 

Myxobohis  transovalis  Gurley  1893 ' (139) 

30(29)    Length  equal  to  or  more  than  breadth  of  spore 31 

31(102)  Length  longer  than  breadth 32 

32(37)     Breadth  of  spore  less  than  half  of  length 33 

33(34)  Extremities  of  spore  equally  pointed;  length  13  to  14.5m,  breadth  6  to  7m,  polar 
capsule  4.5m  long 

Myxobolus  miyairii  Kudo  1919 (155) 

34(33)    Anterior  end  of  spore  attenuated;  posterior  end  rounded 35 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


429]  STUDIES  ON  MYXOSPORIDIA—KUDO  191 

35(36)  Shell  thickened  at  posterior  margin;  spore  12  to  ISy.  long,  4  to  6.8^  broad,  polar 
capsule  5.5  to7juby2.1  to2.5/i 

Myxobolus  anurus  Cohn  1895 (142) 

36(35)     Shell  of  uniform  thickness;  spore  14 .  3n  long,  6.7|x  broad,  polar  capsule  6.5/*  by  2/t 

Myxobolus  funduli  Kudo  1919 (ISl) 

37(32)     Breadth  of  spore  greater  than  half  of  length 38 

38(41)     Length  of  spore  greater  than  20 ft 39 

39(40)  Spore  large;  subcircular  and  anterior  end  flattened  in  front  view;  sutural  edge 
with  markings;  length  38  to  45 ju,  breadth  32  to  38^1,  thickness  28  to35A(, polar 
capsule  15  to  17/i  long 

Myxobolus  magnus  Awerinzew  1913 (ISO) 

40(39)  Spore  small;  extremities  equally  rounded;  length  21/*,  breadth  15/i,  polar  capsule 
6n  long 

Myxobolus  cyprini  Doflein  1898 (143) 

41(38)     Length  of  spore  less  than  20/t 42 

42(43)  Spore  with  a  wide  (2  to  Zn)  membraneous  posterior  process;  length  12/x,  breadth 
10/i,  length  of  polar  capsule  4.5/t 

Myxobolus  cordis  Keysselitz  1908 (148) 

43(42)     Normal  spore  without  appendage 44 

44(49)     Breadth  of  sutural  ridge  one-third  of  thickness  of  spore 45 

45(46)  Length  of  spore  smaller  than  10/t;  subcircular  in  front  view;*length  7  to  8/*, 
breadth  6  to  7/x,  thickness  5/x 

Myxobolus  globosus  Gurley  1893 (139) 

46(45)     Length  of  spore  greater  than  10/* 47 

47(48)  Spore  large;  elliptical  in  front  view;  with  an  intercapsular  appendix,  sutural  edge 
with  markings;  length  16.9  to  21.6/*,  breadth  13  to  16.2/*,  thickness  9/* 

Myxobolus  gigas  Auerbach  1906 (145) 

48(47)  Spore  small;  subcircular  in  front  view;  markings  on  entire  sutural  edge; 
length  10.8  to  11.7/*,  breadth  9.9  to  10.4/*,  thickness  7.2  to  9/* 

Myxobolus  aeglefini  Auerbach  1906 (144) 

49(44)     Sutural  ridge  narrower 50 

50(69)    Spore  with  an  intercapsular  appendix 51 

51(68)     Intercapsular  appendix  triangular 52 

52(57)     Anterior  end  of  spore  attenuated  in  front  view 53 

53(54)     Sutural  edge  without  marking;  length  11  to  12/*,  breadth  9.25  to  10.5/i 

Myxobolus  balleri  Reuss  1906 (147) 

54(53)     Sutural  edge  with  markings 55 

55(56)     Spore  small;  length  8  to  9 . 5/*,  breadth  6  to  7 . 5/*,  thickness  5.5/* 

Myxobolus  exiguus  Thelohan  1895 (1^) 

56(55)  Spore  large;  often  subcircular;  length  1 1 . 5/t  to  12/*,  breadth  7 . 5  to  8/*,  thickness 
5/x 

Myxobolus  obesus  Gurley  1893 (140) 

57(52)    Anterior  end  of  spore  broadly  rounded  in  front  view > 58 

58(59)  Posterior  portion  of  spore  narrower;  polar  capsule  rather  large;  length  11.4  to 
13.5/*,  breadth  9.5  to  U/t,  thickness  8.5  to  9.5/*,  polar  capsule  5.5  to  6/*  long 

Myxobolus  discrepans  Kudo  1919 (156) 

59(58)     Extremities  of  spore  approximately  equal 60 

60(61)  Sutural  edge  without  markings;  spore  10  to  10.5/*  long,  8  to  8.5/*  broad,  polar 
capsule  4.5/*  long 

Myxobolus  squamae  Keysselitz  1908 (147) 

61(60)     Sutural  edge  with  markings 62 

62(65)    Markings  distinct 63 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


192  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [430 

63(64)  Marking  variable  in  number  along  posterior  margin  of  spore;  spore  more  elon- 
gated; length  12  to  12.5a(,  breadth  10  to  IO.Sai,  polar  capsule  5.5  to  6^  long 

Myxobolus  pfeiferi  Thdohan  1895 (133) 

64(63)  Sutural  edge  with  four  markings  around  posterior  margin;  spore  rather  short; 
length  1 1  to  12m,  breadth  9  to  9.5At,  thickness  4.5  to  5m,  polar  capsule  5m  by  2.5m 

Myxobolus  scardinii  Reuss  1906 (14^) 

65(62)    Markings  indistinct 66 

66(67)  Markinp  about  five  at  posterior  margin;  spore  larger  and  shorter;  length  11  to 
12m,  breadth  9.25  to  10m,  thickness  4.5  to  5 .  5m,  polar  capsule  4  to  5m  by  2 .  25m 

Myxobolus  bramae  Reuss  1906 (J'i?) 

67(66)  Markings  many  along  entire  sutural  edge  except  anterior  tip;  spore  smaller, 
longer  and  thicker;  length  9 .  25  to  10m,  breadth  7  to  7 .  25m,  thickness  5  to  5 .  5m, 
polar  capsule  4. 5m  by  2 . 5  to  3m 

Myxobolus  cyprinicola  Reuss  1906 (l^) 

68(51)  Intercapsular  appendix  rounded;  sutural  edge  smooth;  length  11m,  breadth  8m, 
polar  capsule  4  to  6m  long 

Myxobolus  musculi  Keysselitz  1908 (i48) 

69(50)     Spore  without  intercapsular  appendix 70 

70(75)     Length  of  spore  less  than  IOm 71 

71(72)  Spore  very  much  flattened  and  small;  length  6m,  breadth  4 . 2  to  5m,  polar  capsule 
3n  by  2m 

Myxobolus  minutus  Nemeczek  1911 (ISO) 

72(71)    Thickness  of  spore  about  half  of  length 73 

73(74)    Shell  thick;  length  9 .  25  to  IOm,  breadth  7 .  25  to  8 .  25m,  thickness  4  to  5m 

Myxobolus  sandrae  Reuss  1906 (146) 

74(73)    Shell  thin;  spore  8.25  to  9.5m  long,  7.25  to  8.25m  broad,  4.5  to  5.5m  thick 

Myxobolus  volgensis  Reuss  1906 (145) 

75(70)    Length  of  spore  greater  than  10m 76 

76(85)     Extremities  of  spore  approximately  equal 77 

77(80)     Spore  elongated  (breadth:  length  =  l:1.8  or  1:1.4) 78 

78(79)    Spore  larger;  length  14  to  17m,  breadth  8.5m,  thickness  5  to  6m 

Myxobolus  oblongus  Gurley  1893 (139) 

79(78)    Spore  smaller;  length  12  to  15  m,  breadth  9  to  11m 

Myxobolus  ellipsoides  Thdohan  1892 (136) 

80(77)    Spore  shorter  (breadth:  length  =  l:1.3,  1:1.2  or  1:1.1) 81 

81(82)  Sutural  edge  with  markings;  slightly  truncate  at  anterior  end  in  front  view;  spore 
9.5  to  11.5m  long,  8.5  to  9. 5m  broad,  6.5m  thick,  polar  capsule  4. 75m  by  1.5 
to  2m 

Myxobolus  mesenlericus  Kudo  1919 (157) 

82(81)    Sutural  edge  without  markings 83 

83(84)    Polar  capsule  larger;  spore  13  .9m  long,  11m  broad,  8m  thick 

Myxobolus  lintoni  Guriey  1893 (138) 

84(83)    Polar  capsules  smaller;  spore  14 . 5  m  long,  1 1 .  9m  broad,  polar  capsule  6m  by  3 .  7m 

Myxobolus  pleuronectidae  Hahn  1917 (152) 

85(76)     Anterior  end  of  spore  more  attenuated  than  posterior 86 

86(89)     Sutural  edge  with  markings 87 

87(88)  Markings  five  or  six  in  number;  spore  17  to  18m  long,  10  to  13m  broad,  polar  cap- 
sule 7  to  8m 

Myxobolus  permagnus  Wegener  1910 (149) 

88(87)  Markings  sometimes  present;  spore  13  to  17m  lo°gj  8  to  10m  broad,  5  to  7m  thick, 
polar  capsule  6  to  7  m  long 

Myxobolus  carassii  Klokacewa  1914 (150) 

Numben  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


431]  STUDIES  ON  MYXOSPORIDIA—KUDO  193 

89(86)     Sutural  edge  without  markings 90 

90(99)     Anterior  end  of  spore  highly  attenuated 91 

91(96)     Length  of  polar  capsule  equal  to  or  less  than  half  of  that  of  spore 92 

92(93)     Spore:  anterior  end  pointed;  length  12.4  to  13.5m,  breadth  6.5  to  7.5/tt,  thick- 
ness 5jix,  polar  capsule  6  to  7/i  long.   Cysts  of  bright  golden  color 

Myxobolus  aureatus  Ward  1919 (JS4) 

93(22)     Anterior  tip  of  spore  not  pointed 94 

94(95)     Spore  greater  in  thickness  (6 . 5  to  7)u) ,  length  12  to  13  ft,  breadth  8 .  25  to  9/*,  polar 
capsule  6/i  by  2.Sm;  anterior  end  more  rounded 

Myxobolus  physophilus  Reuss  1906 (1^) 

95(94)     Spore  smaller  in  thickness  (5.5/ti),  length  11  to  13^,  breadth  8.25  to9.25A»,  polar 
capsule  6/i  by  2 . 5  to  3m;  anterior  end  less  rounded 

Myxobolus  macrocapsularis  Reuss  1906 (146) 

96(91)     Length  of  polar  capsule  greater  than  half  of  that  of  spore 97 

97(98)     Length  of  polar  capsule  greater  than  two-thirds  of  that  of  spore;  spore  16^  long, 
10  to  11m  broad,  polar  capsule  11m  by  4m 

Myxobolus  capsulalus  Davis  1917 (i52) 

98(97)     Length  of  polar  capsule  less  than  two-thirds  of  that  of  spore;  spore  14  to  16m  long, 
8  to  9m  broad,  5  to  6m  thick,  polar  capsule  8  to  9m  by  2 . 5  to  3  m 

Myxobolus  koi  Kudo  1919 (155) 

99(90)     Anterior  end  of  spore  rounded 100 

100(101)  Cysts:  size  up  to  1.7  mm.  by  0.7  mm.;  parasitic  in  various  tissues  of  host. 
Spore  10  to  12m  long,  9m  broad 

Myxobolus  oviformis  Th6lohan  1892 (137) 

101(100)  Cysts:  0.9  mm.  by  0.02  mm.;  parasitic  in  nervous  system.    Spore  10  to  12m 
long,  8m  broad,  6m  thick,  polar  capsule  6  to  7m  by  2m 

Myxobolus  neurobius  Schuberg  et  Schroder  1905 (144) 

102(31)  Spore  almost  circular  in  front  view 103 

103(104)  Anterior  end  somewhat  attenuated;  sutural  edge  with  four  markings;  spore  9 
to  10m  long  and  broad,  6.5  to  7m  thick,  polar  capsule  6  to  7.5m  by  2.5  to  3m 

Myxobolus  orbicuiatus  Kudo  1919 (1S5) 

104(103)  Spore  regularly  circular  in  front  view lOS 

105(106)  Cysts  large,  up  to  3  mm.  by  1 .5  mm.    Spore  10m  long,  9.8m  broad,  3m  thick, 
polar  capsule  3.8  to  5m  long 

Myxobolus  roiundus  Nemeczek  1911 (149) 

106(105)  Cysts  small,  up  to  0.33  mm.  in  diameter.    Spore  9m  in  diameter 

Myxobolus  sphaeralis  Gurley  1893 (141) 

Incompletely  described  species 

Myxobolus  sp.  Guriey  1894 (142) 

Myxobolus  sp.  Guriey  1894 , (142) 

Myxobolus  sp.  Guriey  1894 (143) 

Myxobolus  sp.  Miyairi  1909 (149) 

Myxobolus  sp.  Wegener  1^10 (149) 

Myxobolus  sp.  Lebzelter  1912 (150) 

Myxobolus  sp.  Southwell  1915 (1^1) 

Myxobolus  sp.  Kudo  1918 • (132) 

Genus    HENNEGUYA    Th61ohan    1892 

1(10)    Parasitic  in  urinary  bladder  or  urinary  tubule  of  kidney  of  host 2 

2(  7)     Shell-valves  striated 3 

Ntunbers  enclosed  ir  pa  rentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


194  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [432 

3(  6)    Length  of  tail  equal  to  two-thirds  of  length  of  main  part  of  spore* 4 

4(  5)  Shell-valves  asymmetrical;  spore  smaller;  total  length  38  to  48/*,  length  of  main 
part  15/i,  breadth  6  to  7.5;^,  polar  capsule  7.5  to  9m  by  3  to  3.5m.  Tropho- 
zoite disporous  and  polysporous 

Henneguya  gasterostei  Parisi  1912 (169) 

S(  4)  Shell-valves  symmetrical;  spore  broader;  length  of  main  part  13.5  to  15ai, 
breadth  8  to  9^,  thickness  6  to  7 .  5m,  polar  capsule  5  to  6m  by  3ft,  length  of  tail 
30  to  40m.    Trophozoite  mictosporous 

Henneguya  mictospora  Kudo  1919 (173) 

6(  3)  Length  of  tail  equal  to  half  of  length  of  main  part  of  spore  and  single(?) ;  length 
20  to  24m,  breadth  5  to  6m,  length  of  polar  capsule  4  to  5m 

Henneguya  media  Thelohan  1892 (161) 

7(  2)     Shell-valves  unstriated 8 

8(  9)  Spore  elongated;  polar  capsule  longer;  posterior  portion  of  main  part  broad;  tail 
wider  and  bifurcated  to  same  direction;  total  length  19.5  to  22.5m,  length  of 
main  part  8.5m,  breadth  6m,  length  of  tail  8  to  8.5m 

Henneguya  Ugeri  Cepdde  1905 (166) 

9(  8)  Spore  oval;  polar  capsule  shorter;  posterior  part  of  main  portion  narrow;  tail 
narrower  and  bifurcated  to  opposite  directions;  length  of  main  part  11.5m, 
breadth  7m,  length  of  tail  9.6m,  polar  capsule  3.5m  by  2.5m 

Henneguya  wisconsinensis  Mavor  et  Strasser  1916 (170) 

10(  1)    Parasitic  in  tissue  of  host 11 

11(28)     Tail  always  appears  as  a  single  process 12 

12(13)     Spore  small;  length  4m,  breadth  2m 

Henneguya  tenuis  Vaney  et  Conte  1901 (166) 

13(12)     Spore  longer  and  larger,  at  least  27m  long 14 

14(15)     Sutural  edge  with  eight  to  ten  markings;  tail  rather  long;  length  of  main  part  10 
to  11  .5m,  breadth  8  to  8 .  75m,  thickness  4  to  5m,  polar  capsule  3  to  4/*  by  2m,  tail 
up  to  17m  long 

Henneguya  brackyura  Ward  1919 (171) 

15(14)     Sutural  edge  without  markings 16 

16(19)    Total  length  of  spore  greater  than  40m 17 

17(18)  Anterior  end  rounded;  polar  capsule  large;  sheU-valves  asymmetrical;  tail  long; 
main  part  10  to  11m  long,  6  to  8m  broad,  4m  thick,  tail  30  to  40m  long.  Cysts 
elongated  and  large  up  to  6  mm.  by  2  mm. 

Henneguya  macrura  Gurley  1894 (164) 

18(17)  Anterior  end  attenuated;  polar  capsule  smaller;  shell-valves  symmetrical;  tail 
shorter;  length  20m,  breadth  8  to  9m,  polar  capsule  8m  by  2  to  3m.  Cysts 
elongated  oval  and  smaller,  1 . 1  mm.  by  0.5  mm. 

Henneguya  creplini  Gurley  1894 (162) 

19(16)    Total  length  of  spore  less  than  40m 20 

20(21)  Tail  about  one- third  of  main  part;  total  length  38m,  niain  part  26m  long,  10  to  11m 
broad,  8m  thick,  polar  capsule  11  to  14m  by  2  to  3m.  Cysts  oval,  numerous  and 
smaU  (130m  by  115m) 

Henneguya  minuta  Cohn  1895 (160) 

21(20)    Tail  about  three-sevenths  of  main  part;  total  length  29  to  40m,  main  part  15  to 

20m  long,  7  to  8m  broad,  polar  capsule  8m  by  2m 22 

22(25)     Cysts  large 23 

*Length  of  main  part  of  spore  denotes  in  all  possible  cases  the  distance  between  the  outer 
anterior  tip  and  the  posterior  margin  of  sporal  cavity;  and  consequently  that  between  the 
latter  and  the  distal  end  of  the  tail  is  the  length  of  the  taiL 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  q>ecies 

named. 


433]  STUDIES  ON  MY XOSPORIDI A— KUDO  195 

23(24)     Cysts  spherical  up  to  6  mm.  in  diameter;  parasitic  in  ovary 

Henneguya  oviperda  (Cohn  1895) (^60) 

24(23)     Cysts  elongated  up  to  2  mm,  by  1 .5  mm.;  parasitic  in  branchia 

Henneguya  psorospermica  Thelohan  1895 (^58) 

25(22)     Cysts  rather  small  or  size  not  observed 26 

26(27)     Elongated  cysts  0.75  mm.  by  0.375  mm.;  parasitic  in  branchia 

Henneguya  texta  (Cohn  1895)  (159) 

27(26)     Parasitic  in  intestinal  wall 

Henneguya  peri-intestinalis  Cep^de  1906 (161) 

28(11)     Tail  composed  of  two  processes 29 

29(30)  Total  length  reaches  87.5  to  110. 5^;  length  of  main  part  10.5  to  15/x,  breadth 
5n,  length  of  polar  capsule  5/i,  length  of  tail  75  to  lOO/x 

Henneguya  gigantea  Nemeczek  1911 (168) 

30(29)    Total  length  of  spore  less  than  S2/j. 31 

31(34)     Sutural  edge  with  markings 32 

32(33)  Tail  longer  and  spore  larger;  anterior  end  more  rounded;  total  length  47 /i.  Cysts 
spherical  and  large,  up  to  6  mm. 

Henneguya  salminicola  Ward  1919 (171) 

33(32)  Tail  shorter  and  spore  smaller;  anterior  end  slightly  more  attenuated;  total 
length  32  ju.    Cysts  lenticular,  up  to  2  mm.  in  length 

Henneguya  niisslini  Schuberg  et  Schroder  1905 (166) 

34(31)     Sutural  edge  without  markings 35 

35(38)     Distal  end  of  tail  thread-like 36 

36(37)  Tail  40  to  50^  in  length;  total  length  50  to  60^,  main  part  8 . 5  to  9 . 5/x  long,  8 . 5  to 
9.5m  broad,  polar  capsule  4.7  to  5. 5m  by  ?>n.  Cysts  small,  up  to  50^  in 
diameter 

Henneguya  neapolitana  Parisi  1912 (170) 

37(36)    Tail  10  to  30/*  in  length;  main  part  12^  long,  8/x  broad 

Henneguya  miyairii  Kudo  1919 (172) 

38(35)    Distal  end  of  tail  tapers  to  a  point  and  not  thread-like 39 

39(40)  Cysts  irregular  in  shape;  size  up  to  2.5  mm.  Spore:  total  length  30  to  40/x, 
main  part  11.5  to  15^  long,  5  to  6.5^  broad,  polar  capsule  6.5  to  8/x  by  2  to 
2.5m,  tail 22  to  28m 

Henneguya  lobosa  (Cohn  1895) (161) 

40(39)    Cysts  spherical  or  oval 41 

41(42)  Anterior  end  of  spore  rounded;  tail  either  single  or  double  processes;  total  length 
55m,  length  of  main  part  10m,  breadth  7m,  length  of  tail  40  to  50m.  Cysts 
spherical  or  oval  up  to  32  mm.  by  16  mm. 

Henneguya  zschokkei  (Gurley  1893) (165) 

42(41)  Anterior  end  attenuated;  spore  large;  main  part  20  to  22m  long,  breadth  8  to 
9m,  6  to  7m  thick,  polar  capsule  10m  by  2  to  ifi,  tail  50  to  60m  long 

Henneguya  acerinae  Schroder  1906 (167) 

Incompletely  described  species 

Henneguya  schizura  (Gurley  1893) (162) 

Henneguya  linearis  (Gurley  1893) (163) 

Henneguya  gurleyi  Kudo  1893 (163) 

Henneguya  monura  (Gurley  1893) (164) 

Henneguya  strongylura  (Gurley  1894) (163) 

Henneguya  kolesnikovi  (Gurley  1894) (164) 

Henneguya  brevis  Thelohan  1895 (162) 

Henneguya  sp.  (Gurley  1894) (165) 

Henneguya  sp.  (Gurley  1894) (165) 

Henneguya  sp.  Nemeczek  1911 (169) 

Numbers  enclosed  in  parentheses  refer  to  the  page  of  the  article  on  which  is  found  the  description  of  the  species 
named. 


196  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [434 


SUMMARY 

1)  All  species  of  Myxosporidia  which  have  been  observed  in  various 
parts  of  the  world,  reaching  237  in  number,  are  recorded  with  figures. 

2)  A  new  classification  of  Myxosporidia  is  proposed  after  discussion 
of  those  of  previous  authors. 

3)  A  complete  list  of  the  specific  names  of  the  hosts  that  harbor  Myxo- 
sporidia, is  given  together  with  the  names  of  the  organ  of  infection  and 
the  localities  from  which  recorded. 

4)  By  study  of  the  geographical  distribution  of  Myxosporidia,  it 
is  shown  that  few  species  are  common  both  to  American  and  European 
waters  or  Asiatic  and  European  waters,  while  the  majority  of  Myxosporidia 
are  localized  in  definite  and  limited  regions. 

5)  The  study  of  the  organal  distribution  of  Myxosporidia  in  the  host, 
shows  that  the  gall-bladder  is  the  organ  most  frequently  invaded  by  the 
parasite.  The  kidney,  branchia  and  urinary  bladder  have  less  chances  of 
being  attacked. 

6)  One  new  genus,  Wardia,  is  established. 

7)  Nine  new  species;  Wardia  ovinocua,  Mitraspora  elongaia,  Chloromy' 
xum  trijugum,  Chloromyxum  catostomi,  Myxidium  americanum,  Myxoholus 
orhiculatus,  Myxoholus  discrepans,  Myxoholus  mesentericus  and  Eenneguya 
mictospora,  are  described  from  fresh-water  fish  collected  in  the  vicinity 
of  Urbana,  HI. 

8)  Six  new  species;  Sphaerospora  carassii,  Myxidium  kagayamai, 
Myxoholus  tnisgurni,  Myxoholus  miyairii,  Myxoholus  koi  and  Eenneguya 
miyairii,  are  recorded  from  fresh-water  fish  of  Nippon. 

9)  One  new  species;  Chloromyxum  wardi,  is  described  from  Alaska. 
This  is  the  second  species  of  Myxosporidia  from  that  part  of  North  America. 

10)  Keys  to  the  genera  and  species  of  known  Myxosporidia  are  in- 
cluded. 


435]  STUDIES  ON  MYXOSPORIDIA—KUDO  197 


APPENDIX:  NEW  MYXOSPORIDIA  FROM  AUSTRALIA 

The  following  six  species  described  by  Johnston  and  Bancroft  did  not 
reach  the  wtiter  until  the  page  proof  was  read.  For  this  reason  they  could 
not  be  put  in  the  text  and  are  recorded  here  separately, 

MYXIDIUM  THERAPON  Johnston  et  Bancroft 
1919    Myxidium  therapon  Johnston  and  Bancroft        1919  :  520-521 

Habitat:  In  the  gall  bladder  of  Therapon  carho  and  Th.  hillii;  Thomson 
River  at  Longreach,  Australia. 

The  parasite  occurred  in  one  specimen  of  the  former  host  fish  and  in 
nine  out  of  thirteen  specimens  of  the  latter.  No  visible  effect  of  the  infec- 
tion on  the  part  of  the  host  fish  was  recognized. 

Vegetative  form:  Body  pale  yellowish  to  green  in  color.  Form(?). 
Size  varies  from  3  to  12mm.  in  diameter.  The  protoplasm  is  differentiated 
into  a  clear  narrow  ectoplasm,  about  10/i  in  width,  and  a  coarsely  grained 
endoplasm.  No  movements  could  be  seen  on  slides;  but  undulations  were 
observed  to  travel  round  the  margin  of  the  trophozoite.    Polysporous. 

Spore:  Spindle-shaped  with  slightly  pointed  extremities.  Polar  cap- 
sules are  more  or  less  rounded.  Shell  with  faintly  marked  longitudinal 
stria tion.  The  sporoplasm  is  binucleated.  Average  dimensions:  length 
9  to  10m,  breadth  4/x,  polar  capsules  2  to  3/i  by  1  to  2/x. 

MYXOSOMA  OGILBYI  Johnston  et  Bancroft 
1919    Myxosoma  ogilbyi  Johnston  and  Bancroft        1919  :  521-522 

Habitat:  In  the  fibrous  tissue  of  the  gill  arch  of  Plectroplites  amhiguus; 
Thomson  River  at  Longreach,  Australia.  Three  out  of  nine  host  specimens 
examined  showed  the  infection. 

Vegetative  form:  The  parasite  forms  white  cysts  usually  close  to  the 
bases  of  the  gill  filaments.  Cysts  are  small  and  rounded,  being  less  than 
1mm.  in  diameter.  The  authors  simply  mention  that  sections  revealed  the 
structure  usually  present  in  a  Myxosporidian  cyst. 

Spore :  Oval  with  pointed  anterior  end.  The  inner  margin  of  the  shell 
is  indented  posteriad.  The  sporoplasm  contains  a  single  nucleus,  but  not 
any  iodinophilous  vacuole.  Average  dimensions:  length  11  to  13ju,  breadth 
6  to  2>ix,  thickness  5/x,  polar  capsules  5  to  6/i  by  2n. 

MYXOBOLUS  PLECTROPLITES  Johnston  et  Bancroft 

1919    Myxobolus  plectroplites     Johnston  and  Bancroft        1919  :  522-523 

Habitat:  In  the  kidney  and  gall-bladder  of  Plectroplites  amhiguus; 
Thomson  River  at  Longreach,  Australia.    The  parasite  was  observed  in 


198  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [436 

four  out  of  nine  host  fish  examined;  in  two  cases  in  the  kidney  only,  in  one 
case  only  in  the  gall-bladder,  and  in  one  instance  in  both  gall-bladder  and 
kidney.  Cysts  were  found  in  the  kidney,  while  only  spores  were  recognized 
in  the  gall-bladder. 

Vegetative  form:  The  cysts  which  could  only  be  detected  in  sections, 
lie  in  the  connective  tissues  of  kidney.  They  are  of  minute  size,  ranging 
somewhat  widely  from  36m  in  diameter  to  144  by  100/i.  According  to  the 
authors  no  definite  structure  could  be  found. 

Spore:  Rounded  oval.  It  bears  quite  a  close  resemblance  to  that  of 
Myxobolus  hylae  (page  153),  which  is  slightly  longer,  and  which  has  a  longer 
polar  filament  than  the  present  form.  The  vacuole,  however,  is  apparently 
not  iodinophilous(?).  Average  dimensions:  length  10  to  12ju,  breadth  7  to 
8/x,  polar  capsules  5  by  2^,  length  of  polar  filament  30  to  40^. 

HENNEGUYA  AUSTRALIS  Johnston  et  Bancroft 
1919    Henneguya  australis         Johnston  and  Bancroft        1919  :  523-524 

Habitat:  In  the  branchiae  of  Plectroplites  amhiguus;  Thomson  River 
at  Longreach,  Australia.  The  parasite  was  detected  in  four  out  of  nine  host 
fish  examined.    The  infection  was  extremely  light  in  all  cases. 

Vegetative  form:  The  parasites  form  cysts.  They  lie  embedded  in  the 
spongy  mass  of  the  gill  filament,  and  in  many  cases  occupy  a  relatively 
large  area  of  the  section.  Cysts  showed  two  layers  in  section;  the  outer- 
most clear  ectoplasm  and  inner  endoplasm  with  developing  spores,  the 
central  portion  of  which  being  filled  with  mature  spores.  The  spores  appear 
to  lie  in  a  definite  manner,  the  long  axis  of  the  spore  commonly  being  at 
right  angles  to  the  boundary  of  the  cyst,  the  anterior  end  of  the  spore 
pointing  outwards. 

Spore :  Elongated  ovoidal.  Anterior  end  pointed,  posterior  end  drawn 
out  into  a  tail.  The  tail  appears  single  when  the  spore  is  removed  from  the 
cyst  but  separates  soon  afterward  into  two  halves  which  usually  diverge 
widely.  Two  polar  capsules  parallel  to  each  other  are  quite  frequently  of 
different  length.  The  sporoplasm  contains  two  nuclei  and  a  small  vacuole 
(iodinophilous?).  Average  dimensions:  length  11  to  IS/x,  breadth  3  to  5/i, 
thickness  3  to  4/z,  polar  capsules  5  to  6^  by  1  to  2jli,  length  of  tail  about  20/Lt. 

HENNEGUYA  GRACILIS  Johnston  et  Bancroft 
1919    Henneguya  gracilis  Johnston  and  Bancroft        1919  :  524-526 

Habitat:  In  the  gill  filament  of  Therapon  hillii;  Thomson  River  at 
Longreach,  Australia.  Out  of  thirteen  specimens  examined,  eight  were 
infected.    Heavy  infection  was  recognized  only  in  one  case. 

Vegetative  form:  The  cyst  is  of  definite,  narrow,  pear-shaped  form,  and 
lie  transversely,  i.e.,  at  right  angles  to  the  long  axis  of  the  gill  filament. 


437]  STUDIES  ON  MYXOSPORIDIA—KUDO  199 

Spore:  The  spore  resembles  Eenneguya  australis,  but  is  slightly 
smaller,  while  the  tail  is  longer  in  proportion.  The  spores  are  arranged 
with  long  axis  parallel  to  that  of  the  cyst.  Average  dimensions:  length 
10  to  14/i,  breadth  2 . 5  to  3^.  thickness  3ju,  polar  capsules  5  to  6/i  by  1  to 
2/1,  length  of  tail  about  20  to  26)u. 

HENNEGUYA  sp.  Johnston  et  Bancroft 
1919    Henneguya  sp,  Johnston  and  Bancroft        1919  :  526 

Habitat:  In  the  branchiae  of  Nematalosa  elongata;  Thomson  River  at 
Longreach,  Australia. 

The  authors  state  that  they  observed  a  number  of  spores  of  a  Henne- 
guya in  the  scrapings  of  the  gill  of  one  of  four  host  fish. 

Vegetative  form:  Undescribed. 

Spore:  Undescribed. 


200  ILLINOIS  BIOLOGICAL  MONOGRAPHS  {438 


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•Original  paper  not  seen. 


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DOFLEIN,  F. 

1898.  Studien  zur  Naturgeschichte  der  Protozoen.    III.  Ueber  Myxosporidien.    Zool. 

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EVERMANN,  B.  W. 

1893.    A  report  upon  investigations  made  in  Texas  in  1891.    Bull.  U.  S.  Fish.  Comm., 
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FiCKERT,  K. 

1895.    Ueber  die  Barbenseuche.    Zeitschr.  Fischer.,  3:209-214. 

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1909.    Ueber  Protozoen  als  Parasiten  der  Fische.     Verb,  zool.-bot.  Ges.  Wien.  59 :  32-48. 
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1888.    On  a  Myxosporidium  infesting  Australian  frogs.    Rep.  Austr.  Ass.  Adv.  Sc,  1 :337. 
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1902.    Sur  les  Myxosporidies  des  Coregones  du  lac  de  Neuchitel.     Arch.  sc.  phys.  nat. 

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1914.  Sur  le  cycle  6volutif  chez  les  myxosporidies.    C.R.  acad.  sci.,  158 :190-192. 

*Original  paper  not  seen. 


441]  STUDIES  ON  MYXOSPORIDIA—KUDO  203 

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1893.  On  the  classification  of  Myxosporidia,  a  group  of  protozoan  parasites  infesting 

fishes.     Bull.  U.  S.  Fish  Comm.,  11 :407-420. 

1894.  The  Myxosporidia,  or  psorosperms  of  fishes,  and  t\ie  epidemic  produced  by  then. 

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1917.  On  the  Sporozoan  parasites  of  the  fishes  of  Woods  Hole  and  vicinity.    I.  Further 

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1909.    Autogamie  bei  Protisten  und  ihre  Bedeutung  fur  das  Befruchtungsproblem. 
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1890.    A  remarkable  flat  worm  in  the  golden  frog.     Proc.  Linn.  Soc.  N.S.W.,  5:661-666. 
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1893.    Ueber  die  Drehkrankheit  der  Regenbogenforelle.    Allg.  Fishereiz.,  18:7-8. 

1895.  Ueber  Fischkrankheiten.     Zeitschr.  Fisch.,  3:179-209;  23  fig. 

1896.  Die  sogenannte  Pockenkrankheit  der  Karpfen.    Allg.  Fischereiz.,  2 1 :2, 186,  etc. 
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*Original  paper  not  seen. 


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HUBER,  J.  C. 

1888.    Ueber  Piesbergens  Fischsporospermien.    Centralbl.  Bakt.  u.  Parasit.,  3:663-664. 
ISHH,  S. 

1915.    Myxosporidiosis  of  Nipponese  eel  (Nipponese).    Jour.  Zool.,  27:372-382;  1  pi. 

1915a.  Lentospora-disease  of  the  eel  (Nipponese).   Jour.  Zool.  27 :471-474;  4  fig. 
Jameson,  P. 

1913.  A  note  on  some  Myxosporidia  collected  at  Monaco.  Bull.  I'inst.  oc6an.,  no.  273. 
JOHKSTON,  T.  H. 

1909.    Exhibit  of  Myxobolus.    Proc.  Roy.  Soc.  N.S.W.,  43:29. 
Johnston,  T.  H.  and  M.  J.  Bancroft. 

1918.  A  parasite,  Myxoholus  hylae  sp.  nov.,  of  the  reproductive  organs  of  the  golden 

swamp  frog,  Hyla  aurea.    Australian  Zoologist,  1 :171-175;  5  textfig.;  1  pi. 

1919.  Some  new  sporozoan  parasites  of  Queensland  freshwater  fish.     Jour.  Proc.  Roy. 

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Johnstone,  J.  and  H.  M.  Woodcock. 

1907.    On  a  Myxosporidian  infection  of  Gadus  esmarkii,  with  a  note  on  the  identification 
of  the  parasite.    Trans.  Biol.  Soc.  Liverpool,  21:204-208;  1  pi. 
Joseph,  H. 

1905.    Chloromyxum  protei  n.  sp.    Zool.  Anz.,  29:450-451. 

1907.  Chloromyxum  protei  n.  sp.,  ein  in  der  Niere  des  Grottenohns  parasitierendes  Myxo- 

sporidium.    Arch.  Protist.,  8:398-412;  1  textfig.,  2  pi. 
Keysselitz,  G. 

1908.  Ueber  durch  Sporozoen  (Mjocosporidien)  hervorgerufene  pathologische  Verander- 

ungen.    Verb.  Ges.  deutsch.  Natur.  u.  Arzte,  Vers.  79:452-453. 
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Klokacewa,  S. 

1914.  Ueber  die  Myxosporidien  der  Karausche.   Zool.  Anz.,  44:182-186;  2  fig. 

KOLESNIKOFF,  N.  F. 

*1886.    O  psorospermiakh  (mikrosporidiakh)  v  muskulature  rib.  Vet.  Vestnik,  Kharkoff, 
5:242-248;  1  pi. 
Kudo,  R. 

1915.  On  a  new  Myxosporidian  from  a  carp  (Nipponese).   Joum.  Zool.,  27 :5l7-523;  2  pi. 

1916.  Contributions  to  the  study  of  parasitic  protozoa.   Ill,  Notes  on  some  Mjrxospori- 

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1917.  Contributions  to  the  study  of  parasitic  protozoa.    II.  Myxobolus  toyamai  nov. 

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1918.  Experiments  on  the  extrusion  of  polar  filaments  of  Cnidosporidian  spores.    Joum. 

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1899.  Sporozoa.     Das  Tierreich.     5  Lief.     180  pp. 
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1897.  Sur  une  Myxosporidie  des  reins  de  la  tortue.    C.R.  soc.  biol.  49:725-726. 

1898.  Surle Myxidiumdanilewskyi.   C.R.  soc.  biol.,  50:27-30;  9 fig. 
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1900.  Sur  une  Myxosporidie  des  voies  biliares  de  I'Hippocampe  (Sphaeromyxa  sahrasesi 

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♦Original  paper  not  seen. 


443]  STUDIES  ON  MYXOSPORIDIA—KUDO  205 

1902.    Sur  la  multiplication  endogene  des  Myxosporidies.  C.R.,  socbiol.,  54:469-472; 

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L£ger,  L. 

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1906.     Sur  la  structure  de  la  parol  sporale  des  Myxosporidies.    C.R.  acad.  sci.,  142: 
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1852.    Parasitismus  und  Parasiten.     Arch,  physiol.  Heilkde,  1 1 :434-436. 
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1851.  Ueber  Psorospermien  und  Gregarinen.  Arch.  Anat.,  Phys.  u.  wiss.  Med.,  18: 
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LlEBERKtJHN,  N. 

1854.  Notice  sur  les  psorospermies.    Bull.  ac.  roy.  Belg.,  21:21-23. 

1855.  Les  psorospermies  des  poissons.  M6m,  cour.  et  m6m.  sav.  etrang.  acad. 
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Linton,  E. 

1891.    On  certain  wart-like  excrescences,  occurring  on  the  short  minnow,  Cyprinodon 

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1912.     Studi  sui  Cnidosporidi.    Pavia,  91  pp.  ;  1  pi. 
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*1888.    Ueber  die  Myxosporidienkrankheit  der  Barben  in  der  Mosel.     Jahresb.  rhein. 
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1896.    Die  Infektion  der  Fische  mit  Myxosporidien.    Die  Natur,  45:284-285;  5  fig. 
1899.    Cystodiscus  immersus.    Verb,  deutsch.    zool.  Ges.,  9:291-293;  1  fig. 
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1889.    Ueber  eine  Myxosporidium  aus  der  Gallenblase  brazilianischer  Batrachier  (Cys- 
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1915.  Studies  on  the  Sporozoa  of  the  fishes  of  the  St.  Andrew's  Region.  Ann.  Rep. 
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1916.  On  the  life-history  of  Ceratomyxa  acadiensis,  a  new  species  of  Myxosporidia  from 
the  eastern  coast  of  Canada.  Proc.  Amer.  Acad.  Arts  and  Sci.,  51 :55 1-574;  3 
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1916a.  Studies  on  the  protozoan  parasites  of  the  fishes  of  the  Georgian  Bay.    Trans. 

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*Original  paper  not  seen. 


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Mayor,  J.  W.,  and  W.  Strasser. 

1916.  On  a  new  Myxosporidian,  Henneguya  wisconsinensis  n.  sp.,  from  the  urinary  blad- 

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Mazzarelli,  G. 

1905.  Sulla  pseudodifterite  degli  Agoni.     Atti  soc.  ital.  sc.  nat.  mUs.  civ.  stor,  nat. 

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1906.  Le  condizioni  della  pesca  nella  provincia  di  Milano.     Riv.  mens,  pesca,  Milano, 

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1885,    Sur  le  r6le  pathog6nique  de  certaines  psorospermies.      Bull.  soc.  zool.  France, 
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1906.     Phenomenes  de  sexualite  chez  Myxobolus  pfeiferi.    C.R.  soc.  biol.,  110:427-428. 
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1908.  Notes  sur  les  Myxosporidies.    Arch.  zool.  exp.,  8JLIII-LXII;  5  figs. 

1909.  Contribution  a  I'^tude  de  la  sexualit6  chez  les  Myxosporidies  et  chez  les  Micro- 

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1917.  Myxoboliasis  tuberosa,  die  Barbenseuche  oder  Beulenkrankheit  der   Barbe. 

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1912.    An  Introduction  to  the  study  of  the  protozoa.    517  pp.;  194  textfig. 
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1890.    Sullo  svilui^o  dei  Myxosporidi.    Boll.  soc.  nat.  Napoli,  4:160-164;  1  pi. 
Miyairi,  K. 

1909.  Guide  to  the  study  of  parasitic  protozoa  (Nipponese).     170  pp.;  8  pi. 
MOROFF,  T. 

1906.    Ueber  die  Gelbsucht  der  Bachforelle.    Osterr.  Fischereiz.  3 :285. 

MiJLLER,  J. 

1841,    Ueber  eine  eigenthiimliche  krankhafte  parasitische  Bildung  mit  specifisch  organi-" 

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mien).    Ber.  preuss.  Akad.  Wiss.  Berlin,  p.  212-221,  246-250,  232  pp.,  4  pi. 
MuLSOw,  K. 

1911.    Ein  neuer  Gehimparasit  des  Karpfens.    Allg.  Fisch.  Zeit.,  36:483-485;  3  fig. 
Nemeczek,  a. 

1911.    Beitrage  zur  Kenntnis  der  Myxo- und  Microsporidien  der  Fische.   Arch.  Protist., 
22:143-169;  19  textfig.,  2  pi. 
Ntifer,  W. 

1905.     Die  Fische  des  Vierwaldstattersees  und  ihre  Parasiten.      Inaug.-Diss,  Basel, 
230  pp.;  21  fig. 
*1905a.  Die  Parasiten  der  Fische  des  Vierwaldstattersees.    Festschr.    50-jahr.    Besteh. 
naturf.  Geselisch.,  Luzem,  pp.  65-232;  4  pi. 
Ohlmacher,  A.P. 

1893.    Myxosporidia  in  the  common  toad,  with  preliminary  observations  on  two  chro- 
mophile  substances  in  their  spores.    Jour.  Amer.  Med.  Assoc,  20:561-567;  1  pi. 
Parisi,  B. 

1910.  Sphaerosporacaudatan.sp.   Zool.  Anz.,  36:253-254;  3  figs. 
*Original  paper  not  seen. 


445]  STUDIES  ON  MYXOSPORIDIA— KUDO  207 

1912.  Primo  contributo  alia  distribusione  geografica  dei  missosporidi  in  Italia.    Atti 

soc.  ital.  sc.  nat.,  50:283-290;  11  fig. 

1913.  Svi)laL  Sphaerospora  caudataVa.ri'&i.    Atti  soc.  ital.  sc.  nat.  51:5-12;  1  pi. 
Perugia,  A. 

*1890.     SuUe  Myxosporidie  dei  pesci  marini.    Boll,  scient.  Pavia,  12:134-139. 
*1891.     Sulle  Myxosporidie  dei  p>esci  marini.    Boll,  scient.  Pavia,  13:22-25;  1  pi. 
Pfeiffer,  L. 

1890.  Die  Protozoen  als  Krankheitserreger.    1  ed.  100  pp.;  34  fig. 

1891.  Die  Protozoen  als  Krankheitserreger.    2  ed.  pp.  216;  91  fig. 

1893.  Die  Parasitismus  des  Epithelcarcinoms  sowie  der  Sarco-,  Micro-  und  Myxo- 

sporidien  in  Muskelgewebe.    Centralbl.  Bakt.  u.  Parasit.,  14:118-130. 
1895.    Die  Protozoen  als  Krankheitserreger.    Nachtrage.    Jena.  pp.  122;  52  fig. 

PlESBERGEN,  F. 

1886.     Eine  neue  Form  von  Fischpsorospermien  aus  Perca  fivviatUis.     Jahresb.  Ver. 
vater.  Naturk.  Wurttemberg,  42:73. 
Plehn,  M. 

1904.  Woher  kommt  die  Drehkrankheit  der  Salmoniden?   Allg.  Fischereiz.,  29 :151-153. 
1904a.  Weiteres  uber  die  Drehkrankheit.    Allg.  Fischereiz.,  29:183-184. 

1905.  Ueber  die  Drehkrankheit  der  Salmoniden  (Lentospora  cerebralis  [Hofer]  Plehn). 

Arch.  Protist.,  5:145-166;  7  textfig.,  1  pi. 

1906.  Die  Drehkrankheit  der  Salmoniden.    Allg.  Fischereiz.,  31 :465-470. 

1906a.  Nochmals  iiber  die  Drehkrankheit  der  Salmoniden.    AUg.  Fischereiz.,  31:516- 

519. 
1910.    Die  pathogene  Bedeutung  der  Myxoboliden  fiir  die  Fische.    Sitz.  Ges.  Morph.  u. 
Physiol.    Munchen,  26:20-27;  3  fig. 
POCHE,  F. 

1913.    Das  System  der  Protozoen.    Arch.  Protist.,  30:125-321;  1  fig. 
Prenant,  a. 

1902.    Striation  et  ciliation  de  la  partie  adherente  du  Myxidium  Ueberkuhni  Biltschli. 

C.R.  soc.  biol,  54:844-846. 
1902a.  Notes  cytologiques.    VII.    Contribution  a  I'etude  de  la  ciliation.    Striation  et 
ciliation  de  la  partie  adherente  du  Myxidium  Ueberkuhni.   Arch.  anat.  micr., 
Paris,  5:212;  7  textfig. 
Railliet,  a. 

1886.    Maladie  des  Barbeaux  causae  par  des  psorospermies.    Bull.  M6m.  soc.  centr. 

med.  v6t.    Paris,  4:134-137. 
1890.    La  maladie  des  Barbeaux  de  la  Mama.    Bull.  soc.  centrale  d'aquic.      Paris, 
2:117-120. 
Remak,  R. 

1852.    Ueber  runde  Blutgerinsel  und  iiber  pigmentgukelhaltige  Zellen.    Arch.  Anat., 
Phys.  u.  wiss.  Med.,  144-146;  1  pi. 
Reuss,  H. 

1906.    Neue  Myxosporidien  von  Susswasserfischen.      Bull.  I'acad.  imp.  sc.  St.-Peters- 
bourg,  25:199-205;  1  pi. 
Ryder,  J.  A. 

1880.    The  psorosperms  found  in  Aphredoderus  sayanus.   Amer.  Nat.,  14:211-212;  2  fiigs. 
Sandeman,  G. 

1894.  On  the  multiple  tumors  in  Plaice  and  Flounders.        Ann.   Rep.    Scott.    Fish. 

Board,  11:391-392;  p.  391-392. 
*Original  paper  not  seen. 


208  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [446 

Schroder,  O. 

1906.    Eine  neue  Myxosporidienart  aus  den  Kiemen  von  Acerina  cernua   {Henneguya 
acerinae  n.  sp.).    Arch.  Protist.,  7:186-196;    1  pi. 
-t4907.    Beitrage  zur  Entwicklungsgeschichte  der  Myxosporidien.   Sphaeromyxa  sabrazesi 
Laveran  et  Mesnil.    Arch.  Protist.,  9:359-381;  3  textfig.,  2  pi. 

1910.  Ueber  die  Anlage  der  Sporocyste  (Pansporoblast)  bei  Sphaeromyxa  sabrazesi 

Laveran  et  Mesnil.    Arch.  Protist,  19:1-5;  10  fig. 

1912.  Myxosporidien.    Prowazek's  Handbuch  Path.  Protoz.,  1^324-336. 
ScHUBERG,  A.  and  O.  Schroder. 

1905.    Myxosporidien  aus  dem  Nervensystem  und  der  Haut  der  Bachforelle  (Myxoholus 
neurobius  n.  sp.  and  Henneguya  nUssUni  n.  sp.).   Arch.  Protist.,  6:45-60;  1  pi. 

SOUTUWKLL,  T. 

1915.    Notes  from  the  Bengal  Fisheries  Laboratory,  Indian  Museum.    On  some  Indian 
Parasites  of  Fish  with  a  note  on  Carcinoma  in  Trout.    Rec.  Ind.  Mus.,  11 :311- 
330;  2  pi. 
Sotn^WEix,  T.  and  B.  Prashad. 

1918.    On  some  Indian  Myxosporidia.   Rec.  Ind.  Mus.,  15 :344r-348;  1  pi. 
Stazzi,  p. 

m    1906.    Psorospermosi  o  myxoboliasi  de  BarbL   Riv.  mens.  pesca.,Milano,  8:14-19;  3  fig. 
Sttrbeck,  G. 

1900.    Eine  neue  Krankheit  beim  Bachsaibling  (Salvelinus  fontinalis).  Allg.  Fischereiz., 
25:367-368. 

1911.  Eine  grosse  Sporencyste  von  Henneguya  zschokkei.    Schweiz.  Fisch.-Zeitg.,  19: 

163-165;  1  fig. 

1913.  Ueber  eine  eigenartige  Form  des  Auftretens  von  Henneguya  zschokkei  Gurley. 

Schweiz.  Fisch.-Zeitg.,  21:30-31. 

1914.  BeitragzurFischpathologie.   lu.  II.  Schweiz.  Fisch.-Zeitg.,  22:296-299. 

TsfeLOHAN,  p. 

1889.    Sur  la  constitution  des  spores  des  Myxosporidies.    C.R.  acad.  sci.,  109 :919-922. 
-x  1890.    Recherches  sur  le  developpement  des  spores  chez  les  Myxosporidies.   C.R.  soc. 
biol.,  2:602-604. 
1890a.  Nouvelles  recherches  sur  les  spores  des  Myxo^xjridies  (structure   et  developpe- 
ment).   C.R.  acad.  sci.,  111:692-695. 
1890b.  Contribution  k I'fitude  des  Mj^xosporidies.   Microgr.,  2:193-213;  1  pi.  Annal. 
1891.    Sur  deux  Sporozoaires  nouveaux,  parasite  des  muscles  des  poissons.    C.R.  soc. 
biol.,  3:27-29. 
fl  fl  «v^  1892.    Observation  sur  les  myxosporidies  et  6ssai  de  classification  de  ces  organismes. 
f'^^  -^  Bull.  soc.  philom.,  4:165-178. 

1892a.  Myxosporidies  de  la  vesicule  biliaire  des  poissons.    C.R.  acad.  scL,  115561-964, 
1091-1094. 

1893.  Alteration  du  tissue  muscvdaire  duhs  k  la  pr&ence  de  Myxosporidies  et  de  microbes 

C.R.  soc.  biol.,  5:267-270. 

1894.  Sur  les  aflSnitfe  r6ciproques  des  Myxosporidies.    C.R.  acad.  sci.,  118:428-430. 

1895.  Recherches  sur  les  Myxosporidies.     Bull.  sc.  France  et  Belg.,  26:100-394;  6  fig., 

3  pi. 
TiojAN,  E. 

1909.    Ein  MjTJobolus  im  Auge  von  Leuciscus  rutilus.   Zool.  Anz.,  34:679-682;  3  figs. 
Tyzzer,  E.  E. 

1900.  Tumors  and  Sporozoa  in  fishes.   Joum.  Boston  Soc.  Med.  Sc,  5:62-68;  1  pi. 
Vaney,  C.  and  A.  Conte, 

1901.  Sur  deux  nouveaux  Sporozoaires  dndospores  parasites  de  VAcerina  cernua  Cuv. 

Aim.  Soc.  Linn.  Lyon,  47:  103-106;  4  fig. 


447]  STUDIES  ON  MYXOSPORJDIA—KUDO  209 

Ward,  H.  B. 

1919.    Notes  on  North  American  Mjrxosporidia.    Journ.  Parasit.,  6:49-64,  1  pi, 
Wasieiewsky,  von 

1896.    Sporozoenkunde.    Jena.    162  pp.;  Ill  textfig. 
Wegener,  G. 

1910.    Die  Ektoparasiten  der  Fische  Ostpreussens.    Inaugur.-Dissert.  Konigsberg,  p. 
72^0;  4  figs. 
Weltnes,  W. 

1892.  Ueber  Myxosporidien  Sporen  in  den  Eiem  von  Esox  lucius.    Sitz.  Ges.  nat.  Fr. 

Berlin,  p.  7-41;  16  fig. 
Whinery,  J.  B. 

1893.  Some  additional  notes  on  a  M3Tcosporidian  infection  in  the  common  toad.   N.  Y. 

Med.  Journ.,  18:660H562;  1  fig. 
Woodcock,  H.  M. 

1904.    On  M3Tcosporidia  in  flat-fish.   Trans.  Biol.  Soc.  Liverpool,  18:46-62;  1  pi. 
1907.    On  a  Myxosporidian  infection  of  Gadus  estnarkii.    With  Johnstone. 

ZSCHOKKE,  F. 

1884.    Psorospermies  de  Coregonus  fera.    Arch.  biol.  gand,  Leipzig  et  Paris,  5:234-235; 

Ipl. 
1898.    Die  Myxosporidien  der  Gattimg  Coregonus.   Centralbl.  Bakt.  u.  Parasit.,  Abt.  L 

Orig.,  23:602-607,  646-655,  699-703;  4  fig. 
1898a.  Die  M3TJOsporidien  in  der  Muskulature  der  Gattung  Coregonus.    Zool.  Anz., 

21:213-214. 
1898b.  Myxobolus  bicaudatus  n.  sp.,  ein  Parasit  der  Coregoniden  des  Vierwaldstattersees. 

Mitt.  Naturf.  Ges.,  Luzem,  p.  205-217. 
1900.    Myxobolus  psorospermicus  Th61ohan  im  Vierwaldstattersee.    Mitt.  Naturf.  Ges. 

Luzem,  p.  441-442. 
ZscHOKKE,  F,  and  A.  Heitz. 

1914.    Entoparasiten  aus  Salmoniden  von  Kamtschatka.     Revue  Suisse  zool.,  22:195- 

256. 


210  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [448 


GENERAL  EXPLANATION  OF  FIGURES 

For  the  type  species  of  each  genus,  both  the  vegetative  form  and  the 
spore  are  illustrated.  For  the  other  species,  except  those  which  are  new, 
figures  of  the  spore  are  given,  unless  the  vegetative  forms  are  different 
from  those  of  the  type  species  or  the  species  were  reported  in  papers  which 
seem  to  be  of  less  universal  distribution. 

The  original  drawings  were  made  with  the  Abbe  drawing  apparatus. 
The  combinations  used  were  Zeiss  apochromatic  objectives  16,  8,  3,  and 
homogeneous  oil  immersion  2  mm.  with  compensation  oculars  2,  4,  6,  8, 
12  and  18.  All  the  other  drawings  were  copied  from  the  original  figures  of 
the  respective  observers,  an  exact  citation  of  which  is  given  in  each  case, 
and  were  also  made  with  the  same  drawing  apparatus  on  the  same  scale 
except  that  a  few  figures  were  enlarged  among  those  that  were  taken 
from  other  authors. 

Magnifications  were  also  calculated  and  given  for  those  quoted  figures, 
for  which  the  respective  authors  failed  to  mention  the  scale  at  which  the 
drawings  were  made. 


44^  STUDIES  ON  MYXOSPORIDIA— KUDO  211 


PLATE  I 


212  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [450 


EXPLANATION  OF  PLATE 
Figs.  1  to  5.    Leptotkeca  agUis. 

Fig.    1.    A  typical  trophozoite  in  vivo.    After  Thfilohan  (1895,  Fig.  29);  X750.  * 
Fig.    2.    A  young  form.    After  Th^lohan  (1895,  Fig.  31).     X750. 
Fig.    3.    A  trophozoite  in  motion.    After  Doflein  (1898,  Fig.  5). 
Fig.    4.    A  trophozoite  in  contracted  condition.    After  Doflein  (1898,  Fig.  7). 
Fig.    5.    A  fresh  spore.    After  Th61ohan  (1895,  Fig.  30).     X1500. 
Fig.    6.    A  fresh  mature  spore  of  Leptotkeca  dongata.    After  Thdohan  (1895,  Fig.  38). 

X1500. 
Fig.    7.    A  fresh  mature  spore  of  Z^/><o//(«ca  ^orra.   After  Thdohan  (1895,  Fig,  25).     X1500. 
Fig.    8.    A  fresh  spore  of  Leptotkeca  perlata.     After  Balbiani  (1884,  Fig.  40). 
Fig.    9.    A  spon  oi  Leptotkeca  macros pora.    After  Auerbach  (1909,  Fig.  2a).     X1350. 
Fig.  10.    A  spore  of  Leptotkeca  informis,  preserved  in  formol.    After  Auerbach  (1910b,  Fig. 

la).     X  about  2000. 
Fig.  11.    A  spore  of  Leptotkeca  longipes,  preserved  in  formol.    After  Auerbach  (1910b,  Fig. 

Id).     X  about  2200. 
Fig.  12.    AiresiispoitoiLeptotkecafusiformis.    After  Davis  (1917,  Fig.  1).     X1500. 
Fig.  13.    Airesh  spore  ol  Leptotkeca  scissura.    After  Davis  (1917,  Fig.  8).     X1500. 
Fig.  14.    A  fresh  spore  of  Leptotkeca  lohosa.    After  Davis  (1917,  Fig.  11).     X1500. 
Fig.  15.    A  fresh  spore  of  Leptotkeca  glomerosa.    After  Davis  (1917,  Fig.  13).     X1500. 
Fig.  16,    A    trophozoite  of  Leptotkeca  sp.    After  Awerinzew  (1908,  PI.  2,  Fig,  14).    1/12 

and  comp,  oc,  12. 
Fig.  17.    Another  trophozoite  of  the  same.    After  Awerinzew  (1908,  PL  2,  Fig.  17).    1/12 

and  comp.  oc,  12, 
Figs.  18  to  20.     Ceratomyxa  arcuata.    After  Th^lohan  (1895). 
Figs.  18  and  19.    Typical  young  form.    After  Thflohan  (1895,  Figs.  16  and  17). 
Fig.  20.    A  trophozoite  with  two  spores.    After  Thelohan  (1895,  Fig.  18). 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


VOLUME  V 


KUDO 


9     ^<:=f==:^        ■"       "~<=i=^       14 
STUDIES  ON  MYXOSPORIDIA 


PLATE  I 


4S1J  STUDIES  ON  MYXOSPORIDIA—KUDO  213 


PLATE  II 


214  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [452 


•  EXPLANATION  OF  PLATE 

Fig.  2L    A  sporulating  trophozoite  of  Ceratomyxa  arcuata.   After  Parisi  (1912,  Fig.  6a). 
Fig.  22.    A  spore  of  Ceratomyxa  arcuata  treated  with  KOH.   After  Th6lohan  (1895,  Fig.  19). 

X1500. 
Figs.  23  and  24.    Ceratomyxa  sphaendosa.   After  Thdlohan. 
Fig.  23.    A  part  of  the  tropzoite  (1895,  Fig.  2).     X750. 
Fig.  24.    A  fresh  spore  (1895,  Fig.  3).     X750. 

Fig.  25.    A  fresh  spore  of  Ceratomyxa  globulifera.    After  Th^lohan  (1895,  Fig.  43).    X 1500. 
Fig.  26.    An  adult  trophozoite  of  Ceratomyxa  appendiculata.    After  Thfilohan  (1895,  Fig.  4) . 

X  about  400. 
Fig.  27.    A  spore  of  Ceratomyxa  truncata.    After  Th^lohan  (1895,  Fig.  51).     X1500. 
Fig.  28.    A  spoK  oi  Ceratomyxa  reticularis.    After  Th61ohan  (1895,  Fig.  27).     X1500. 
Fig.  29.    A  siporeoi  Ceratomyxa  inaequclis.    After  Doflein  (1898,  Fig.  10).     X1250. 
Figs.  30  and  31.    Ceratomyxa  linospora.    After  Doflem  (1898).     X  about  1900. 
Fig.  30.    A  spore  (1898,  Fig.  39). 
Fig.  31.    A  trophozoite  with  spores  (1898,  Fig.  43). 
Figs.  32  and  33.    Ceratomyxa  ramosa.    After  Awerinzew  (1908). 
Fig.  32.    A  trophozoite  (1908,  PI.  2,  Fig.  20).    Zeiss  obj.  D  and  comp.  oc.  4. 
Fig.  33.    A  spore  (1908,  PI.  2,  Fig.  19).    Zeiss  obj.  E  and  comp.  oc.  4. 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


VOLUME  V 


KUDO 


STUDIES  ON  MYXOSPORIDIA 


PLATE  II 


453]  STUDIES  ON  MYXOSPORIDIA— KUDO  215 


PLATE  III 


216  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [454 


EXPLANATION  OF  PLATE 

Figs.  34  to  39.    Ceratomyxa  drepanopsettae.    Awerinzew  (1908). 

Fig.  34  and  35.    Trophozoites  (1908,  PL  2,  Figs.  7  and  9).    Obj.  D  and  oc.  4. 

Fig.  36.    The  part  of  a  trophozoite  attached  to  the  epithelium  of  the  gall-bladder  of  the  host 

1908,  PL  2,  Fig.  10).    Obj.  E  and  oc.  4. 
Figs.  37  to  39.    Spores  (1908,  PL  1,  Figs.  2,  3  and  1).    Obj.  D  and  oc.  4. 
Figs.  40  and  41.    Two  different  views  of  the  spore  of  Ceratomyxa  tylosuri.    After  Awerinzew 

(1913a,  Fig.  1).     X  about  350.  4 

Figs.  42  and  43.    Ceratomyxa{?)  spari.    After  Awerinzew  (1913a,  Fig.  2).  * 

Fig.  42.    A  trophozoite. 
Fig.  43.    A  spore.     X  about  345. 
Figs.  44  to  47.    Spores  of  Ceratomyxa  acadiensis.    After  Mavor  (1916).    Figs.  44  and  45 

(1916,  Fig.  B).     X270.    Fig.  46  (1916,  Fig.  A)     X1800.    Fig.  47  (1916,  Fig.  40)  X2950. 
Fig.  48.    A  spore  of  Ceratomyxa  coris.    After  Georg6vitch  (1916a,  Fig.  1). 
Fig.  49.    A  spore  of  Ceratomyxa  herouardi.    After  Georg6vitch  (1917,  Fig.  1). 
Fig.  50.    A  spore  of  Ceratomyxa  mesospora.    After  Davis  (1917,  Fig.  15).     X1500. 
Fig.  51.    A  spore  of  Ceratomyxa  sphairopkora.    After  Davis  (1917,  Fig.  23).     X950. 
Figs.  52  and  53.    Spores  oi  Ceratomyxa  taenia.    After  Davis  (1917,  Figs.  26  and  25).    X700. 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


VOLUME  V 


KUDO 


STUDIES  ON  MYXOSPORIDIA 


PLATE  III 


4SS]  STUDIES  ON  MYXOSPORIDIA—KUDO  217 


PLATE  IV 


218  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [456 


EXPLANATION  OF  PLATE 

Fig.  54.    A  spore  of  Ceratomyxa  aUenuata.    After  Davis  (1917,  Fig.  28).     X950. 

Figs.  55  and  56.    Spores  of  Ceratomyxa  recurvata.    After  Davis  (1917,  Figs.  32  and  33). 

X1500. 
Figs.  57  to  60.    Spores  of  Ceratomyxa  lunata.    After  Davis  (1917,  Figs.  34  to  37).     X 1500. 
Fig.  61.    A  spore  oi  Ceratomyxa  abbreviata.    After  Davis  (1917,  Fig.  41).     X1500. 
Fig.  62.    A  spore  of  Ceratomyxa  flagdlifera.    After  Davis  (1917,  Fig.  42).     X1500. 
Fig.  63.    A  spore  oi  Ceratomyxa  agglomerata.    After  Davis  (1917,  Fig.  45).     X1500. 
Fig.  64.    A  spore  oi  Ceratomyxa  amorpha.    After  Davis  (1917,  Fig.  47).     X1500. 
Figs.  65  to  67.    Spores  of  Ceratomyxa  monospora.    After  Davis  (1917,  Figs.  57,  56  and  55). 

X1500. 
Figs.  68  and  69.    Spores  of  Ceratomyxa  streptospora.    After  Davis  (1917,  Figs.  59  and  60). 

X1500. 
Fig.  70.    A  spore  oi  Ceratomyxa  aggregata.    After  Davis  (1917,  Fig.  63).     X1400, 
Fig.  71.    A  spore  of  Ceratomyxa  undulata.    After  Davis  (1917,  Fig,  66),     X1500. 
Figs.  72  and  73.    Spores  of  Ceratomyxa  navicularia.    After  Davis  (1917,  Figs.  69  and  68). 

X1500. 
Fig.  74.    A  spore  oi  Ceratomyxa  spinosa.    After  Davis  (1917,  Fig.  72).     X1500. 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


VOLUME  V 


KUDO 


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EXPLANATION  OF  PLATE 

Figs.  75  to  80.    Myxoproteus  anibiguus.    After  Doflem  (1898). 

Figs,  75  and  76.    Trophozoites  of  typical  forms  (1898,  Figs.  12  and  52). 

Figs.  77  and  78.    Young  trophozoites  produced  by  budding  (1898,  Figs.  55  and  56). 

Figs.  79  and  80.    Spores  (1898,  Figs.  54  and  64).     X  about  800  and  1080. 

Figs.  81  to  83.    Spores  of  Myxoproteus  cordiformis.    After  Davis  (1917,  Figs.  79,  80  and  78). 

X1500. 
Fig.  84.    ks^itoi  Myxoproteus  cornutus.    After  Davis  (1917,  Fig.  85).     X1400. 
Figs.  85  to  95.    Wardia  ovitwcua.    Original. 
Fig.  85.    A  portion  of  the  cross-section  thru  an  infected  ovary  of  Lepomis  humilis,  showing 

the  parasite  in  one  ovum  and  the  hypertrophied  nurse  cells  and  several  connective  tissue 

layers.     X  160. 
Fig.  86.    A  portion  of  the  cross-section  of  a  cyst.     X640. 
Figs.  87  to  89.    Three  diflEerent  views  of  fresh  spore.     X2000. 
Figs.  90  and  91.    Stained  spores.     X1700. 
Fig.  92.    A  spore  mechanically  pressed  and  stained  with  Giemsa's  mixature,  showing  the 

escaping  polar  capsules  without  extruding  polar  filament,  and  the  sp>oroplasm.     X1700. 
Figs.  93  to  95.    Front  and  lateral  views  of  the  shell  valves,  exhibiting  the  network-like  fine 

ridges  on  the  surface  and  the  posterior  processes.     X1700. 
Figs.  96  and  97.    Spores  of  Wardia  ohlmacheri.    After  Ohlmacher  (1893,  Figs.  4a  and  2). 

2mm.  and  oc.  4. 


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EXPLANATION  OF  PLATE 
Figs.  98  to  104.    Mitraspora  cyprini. 
Figs.  98  and  99.    Trophozoites  from  the  ureter  of  Cyprinus  carpio  in  vivo.    Original.  X 

about  1500. 
Figs.  100  and  101.    Different  views  of  fresh  spores.    Original.     X  about  2000. 
Figs.  102  to  104.     Spores.    After  Fujita  (1912,  Figs,  la  to  Ic). 
Figs.  105  to  107.    Mitraspora  caudaia.    After  Parisi  (1910  and  1913). 
Fig.  105.    A  trophozoite  (1910,  Fig.  1). 
Figs.  106  and  107.    Front  and  lateral  views  of  the  spore  (1913,  Fig.  20  and  1910,  Fig.  2). 

X  about  1600. 
Figs.  108  to  113.     Chloromyxum  leydigi. 

Figs.  108  and  109.    Trophozoites.    After  Th^lohan  (1895,  Figs.  7  and  6).     X750. 
Figs.  110  and  111.    Trophozoites  in  division.    After  Doflein  (1898,  Figs.  57  and  58). 
Figs.  112  and  113.    Different  views  of  spores.    After  Th61ohan  (1895,  Figs.  10  and  9). 

X1500. 
Fig.  114.    A  spore  of  Chloromyxum  catidatum.    After  Thdohan  (1895,  Fig.  36).     X1500. 
Figs.  115  and  116.    Different  views  of  the  spore  of  Chloromyxum  quadratum.    After  Th61ohan 

(1895,  Figs.  100a  and  100b).     X1500. 
Fig.  117.    A  spore  of  Chloromyxum  quadratum  treated  with  nitric  acid.    After  Th6Iohan 

(1895,  Fig.  100c).     X1500. 
Fig.  118.    A  spoie  oi  Chloromyxum  fluviaiile.    After  Th61ohan  (1892,  Fig.  2).     X1500. 
Figs.  119  and  120.    Different  views  of  the  spore  of  Chloromyxum  mucronatum.    After  Lieber- 

kiihn  from  Guriey  (1894,  PI.  39,  Fig.  5).     X  about  1750. 
Figs.  121  and  122.    The  same  after  Balbiani  (1884,  Fig.  41).     X  about  1200. 
Figs.  123  and  125.    Spores  of  Chloromyxum  diploxys.    After  Balbiani  from  Guriey  (1894 

PI.  42,  Figs.  13a,  13b  and  13c). 


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EXPLANATION  OF  PLATE 

Fig.  126,    Trophozoite  of  Chloromyxum  truUae.    After  L^ger  (1906,  Fig.  4).     XlOOO. 

Figs.  127  and  128.  Trophozoites  of  Chloromyxum  cristatum.  After  L6ger  (1906,  Fig.  7). 
XlOOO. 

Figs.  129  to  133.    Chloromyxum  dubium.    After  Auerbach  (1908). 

Figs.  129  and  130.    Trophozoites  (1908,  Figs.  2  and  1). 

Figs.  131  and  132.    Spores  (1908,  Figs.  3  and  4).     X  about  2250. 

Fig.  133.    A  stained  young  spore  (1908,  Fig.  5). 

Fig.  134.  Trophozoite  of  Chloromyxum  sp.  After  Awerinzew  (1908,  PI.  2:  Fig.  12).  Objec- 
tive D  and  ocular  4. 

Fig.  135.    Chloromyxum  koi.    After  Fujita  (1913).     X  about  800. 

Figs.  136  to  138.    Chloromyxum  magnum.    After  Awerinzew  (1913a,  Fig.  4). 

Figs.  136  and  137.    Trophozoites. 

Fig.  138.    A  spore.     X  about  320. 

Figs.  139  and  140.  Spores  oi  Chloromyxum  funduli.  After  Hahn  (1915,  Figs.  34  and  33). 
X2000. 

Figs.  141  to  146.    Chloromyxum  misgurni.    After  Kudo  (1916).    X1750. 

Figs.  141  and  142.    Trophozoites  (1916,  Figs.  3f  and  3g) 

Figs.  143  to  145.    Different  views  of  fresh  spore  (1916,  Figs.  3a,  3c  and  3b). 

Fig.  146.    A  spore  treated  with  potassium  hydrate  (1916,  Fig.  3e). 

Figs.  147  to  152.    Chloromyxum  fujitai.    After  Kudo  (1916).     X1750. 

Fig.  147.    A  trophozoite  (1916,  Fig.  4a), 

Fig.  148.    A  fresh  spore  (1916,  Fig.  4e). 

Fig.  149.    A  spore  stained  with  Giemsa's  mixture  (1916,  Fig.  4g). 

Figs.  150  to  152.  Two  surface  views  and  an  optical  cross-section  of  a  stained  spore,  showing 
the  ridges  on  the  shell  valves  (1916,  Figs.  4b,  4d  and  4c), 

Figs.  153  to  156.    Spores  of  Chloromyxum  clupeidae. 

Figs.  153  to  155.    Fresh  spares.    After  Linton  (1901,  Fig.  3).    "Variously  magnified." 

Fig.  156.    A  spore.    After  Hahn  (1917b,  Fig.  10).     X1650. 

Figs.  157  and  158.  Two  views  of  Chloromyxum  granulosum.  After  Davis  (1917,  Figs.  137  and 
138).     X1500. 


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Figs.  159  to  182.    Chloromyxum  trijugum.    Original. 

Figs.  159  to  174.    Trophozoites  of  various  form  and  age. 

Figs.  159  and  160.    Trophozoites  of  medium  size.     X640. 

Fig.  161,    A  trophozoite,  ten  minutes  after  it  was  removed  from  the  host.     X1700. 

Figs.  162  and  163.  The  movements  of  pseudopodia  of  the  same  specimen  in  ten  minutes. 
X1700. 

Fig.  164.    The  same  specimen  after  thirty  minutes.     X1700. 

Figs.  165  to  167.  A  trophozoite,  showing  the  change  of  pseudopodia  in  five  and  ten  minutes. 
X1700. 

Figs.  168  to  172.  Small  trophozoite  with  different  nimibers  of  the  nuclei.  Fig.  172  is  prob- 
ably a  disporous  form.     X2350. 

Fig.  173.    A  monosporous  trophozoite  with  a  young  spore.     X2350. 

Fig.  174.    A  young  spore.     X2350. 

Figs.  175  to  177.    Different  views  of  fresh  spores.     X1700. 

Fig.  178.    A  Giemsa  stained  spore.     X1700. 

Figs  179  and  180.    Side  views  of  the  valves  showing  the  ridges  by  Giemsa  staining.     X1700. 

Fig.  181.  A  spore  treated  with  potassium  hydrate  solution,  and  stained  with  Giemsa  solu- 
tion.    X1700. 

Fig.  182,  A  spore  from  which  the  sporoplasm  is  leaving  the  shelL  From  the  Giemsa  smear 
of  the  infected  bUe.     X 1700. 


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Fig.  183  to  186.    Sphaerospora  divergens. 

Fig.  183.    Trophozoite.    After  TWlohan  (1895,  Fig.  12).     X7S0. 

Figs.  184  and  185.    Two  views  of  spore.    After  Auerbach  (1912,  PI.  5,  Fig.  4).     X  about 

1500. 
Fig.  186.    A  spore  treated  with  nitric  acid.    After  Thflohan  (1895,  Fig.  13).     X1500. 
Figs.  187  and  88.    Spores  of  Sphaerospora  elegans.    After  Th61ohan  (1890b,  Fig.  1).     X 

about  1000. 
Fig.  189.    A  spore  of  Sphaerospora  rostrata.    After  Thdiohan  (1895,  Fig.  93).     Xabout  1635. 
Figs.  190  to  192.    Spores  of  Sphaerospora  masovica.    After  Cohn  (1902,  Fig.  3).     X 1000. 
Fig.  192.    Spore  with  extruded  polar  filaments  and  "starren  Ffiden"  by  warming. 
Figs.  193  and  194.    Spores  of  Sphaerospora  platessae.    After  Woodcock  (1904,  Fig.  7d). 

X900. 
Figs.  195  to  197.    Spores  of  Sphaerospora  angulata.    After  Fujita  (1912,  Fig.  3).     X  about 

2800. 
Figs.  198  and  199.    Spores  of  Sphaerospora  polymorpha.    After  Davis  (1917,  Figs.  91  and  92) 

X1500. 
Figs.  200  to  204.    Sphaerospora  carassii.    OriginaL 
Fig.  200.    A  trophozoite.     X2250. 
Figs.  201  to  203.    Different  views  of  spores.     XI 800. 
Fig.  204.    A  young  spore.     X2250. 

Figs.  205  to  209.    Sinuolinea  dimorpha.    After  Davis  (1916). 
Fig.  205.    A  fresh  trophozoite  (1916,  Fig.  1).     X1400. 
Figs.  206  and  207.    Trophozoites  with  erythrocytes  in  different  stages  of  disintegratioa 

(1916,  Figs.  2  and  57).     X640. 
Fig.  208.    A  stained  disporous  trophozoite  (1916,  Fig.  41). 
Fig.  209.    A  stained  genmiule  (1916,  Fig,  72). 


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EXPLANATION  OF  PLATE 

Figs.  210  to  213.    Sinuolinea  dimorpha.    After  Davis  (1916  and  1917). 

Fig.  210.    A  living  trophozoite  (1916,  Fig.  56).     X640. 

Fig.  211.    Alivingtrophozoitefromwhichagenunuleis  just  escaping  (1916,  Fig.  60).     X640L 

Figs.  212  and  213.    Spores  (1917,  Figs.  99  and  100).     X1400. 

Figs.  214  to  216.    Spores  of  Sinuolinea  capsularis.    After  Davis  (1917,  Figs.  105  to  107). 

X1500. 
Figs.  217  and  218.    Spores  of  Sinuolinea  arborescens.    After  Davis  (1917,  Figs.  109  to  110). 

X1500. 
Fig.  219.    Spore  oi  Sinuolinea  opacita.    After  Davis  (1917,  Fig.  112).     X1500, 
Fig.  220.    Spore  ot  Sinuolinea  brackiophora.    After  Davis  (1917,  Fig.  113).     XlSOO. 
Figs.  221  to  224.    Myxidium  lieberkukni.    After  Butschli  (1881  and  1882). 
Fig.  221.    A  trophozoite  (1882,  PI.  ^,  Fig.  12).     X  about  60. 
Fig.  222.    A  trophozoite  (1882,  PI.  38,  Fig.  13).     Xl60. 
Figs.  223  and  224.    Trophozoites  (1881,  Figs.  27  and  26). 


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Figs.  225  to  240.    Myxidium  lieherkUhni. 

Fig.  225.    A  trophozoite.    After  LieberkOhn  from  Gurley  (1894,  PI.  43,  Fig.  la).     X330. 

Figs.  226  and  227.    Trophozoites.    After  Btitschli  (1881,  Figs.  25  and  31). 

Figs.  228  to  230.    Stamed  trophozoites.    After  Thflohan  (1895,  Figs.  44, 46  and  45).     X7S0. 

Fig.  231.    A  trophozoite  forming  daughter  individuals  by  budding.    After  Cohn  (1895 

Fig.  5). 
Fig.  232.    Four  figures  showing  the  separation  of  a  bud.    After  Cohn  (1895,  Figs.  6a,  6b, 

6d  and  6e). 
Fig.  233. 1^  A  cross-section  of  a  trophozoite,  showing  the  ectoplasm,  mesoplasm  and  endo- 

plasm.    After  Cohn  (1895,  Fig.  2). 
Fig.  234.    A  trophozoite.    After  Laveran  and  Mesnil  (1902,  Fig.  3).     X500. 
Fig.  235.    A  part  of  a  trophozoite.    After  Laveran  and  Mesnil  (1902,  Fig.  3),     X800. 
Figs.  236  and  237.    Young  trophozoites  undergoing  division.    After  Laveran  and  Mesnil 

(1902,  Figs.  4  and  5).     X 1000. 
Fig.  238.    An  isolated  spore.    After  BUtschli  (1881,  Fig.  32).     X  about  2500. 
Figs.  239  and  240,    Fresh  and  stained  spores.    After  Th6Iohan  (1895,  Figs.  47  and  48). 

X1500. 
Figs.  241  to  251.    Myxidium  incurvatum. 

Fig.  241.    A  monosporous  trophozoite.    After  Parisi  (1912,  Fig.  1). 
Fig.  242.    A  spore.    After  Th61ohan  (1895,  Fig.  54).     X1500. 
Figs.  243  and  244.    Different  views  of  spore.    After  Parisi  (1912,  Fig.  1). 
Fig.  245,    A  young  spore.    After  Georgdvitch  (1916,  Fig.  11). 
Figs.  246  to  248.    Spores.    After  Georg6vitch  (1916,  Figs.  10,  9  and  8). 
Figs.  249  to  251.    Spores.    After  Davis  (1917,  Figs.  119  to  121).     X1500. 
Fig.*252.    Trophozoite  of  Myxidium  sphaericum.    After  Th61ohan  (1895,  Fig.  28).     XlSOO. 
Fig.*253.    A  spore  of  Myxidium  histophUum.    After  Th6Iohan  (1895,  Fig.  49).     X 1500. 
Fig,^254.    A  spore  of  Myxidium  sp.    After  Leydig  from  Gurley  (1894,  PI.  47,  Fig.  6). 


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Figs.  255  to  257.    Myxddium  danilewskyi.    After  Laveran  (1898). 

Figs.  255  and  256.    Longitudinal  and  transverse  sections  thru  renal  tubules,  showing  the 

trophozoites  (1898,  Figs.  1  and  2).     Fig.  255.     X350. 
Fig.  257.     Spores  (1898,  Figs.  4,  5  and  6).     X800. 

Fig.  258.     Spores  of  Myxidium  giganteum.    After  Doflein  (1898,  Fig.  48).     X  about  500. 
Figs.  259  to  261.     Spores  of  Myxidium  giardi.    After  CepSde  (1906a,  Figs.  1,  3  and  2). 

X2000. 
Figs.  262  to  265.     Spwres  of  Myxidium  pfeifferi.    After  Auerbach  (1908,  Figs.  6  and  7). 

X  about  2000,  except  Fig.  265. 
Fig.  266.    A  spore  of  Myxidium  inflatum.    After  Auerbach  (1909,  Fig.  3a).     X  about  1500. 
Fig.  267.    A  spore  of  Myxidium  bergense.    After  Auerbach  (1910a,  Fig.  57).     X  about  1820. 
Fig.  268.    A  spore  of  Myxidium  procerum.    After  Auerbach  (1910a,  Fig.  58).     X  about  2000. 
Figs.  269  to  271.     Spores  of  Myxidium  mackiei.    After  Bosanquet  (1910,  Fig.  12).     X 

about  1250. 
Figs.  272  and  273.     Spores  of  Myxidium  macrocapsidare.    After  Auerbach  (1910d,  Figs,  la 

and  lb).     X3000. 
Figs.  274  to  276.    Myxidium  sp.    After  Awerinzew  (1908  and  1911). 
Fig.  274.    A  monosporous  trophozoite  (1911,  Fig.  C). 
Fig.  275.      A  disprous  trophozoite  (1908,  PI.  2,  Fig.  6).    Obj.  E  and  oc.  4. 
Fig.  276.    A  spore  (1908,  PI.  la.  Fig.  17).     X  about  1000. 
Figs.  277  and  278.     Spores  of  Myxidium  depressum.    After  Parisi  (1912,  Figs.  2a  and  2b). 

X  about  1600. 
Figs.  279  and  280.     Spores  of  Myxidium  oviforme.    After  Parisi  (1912,  Fig.  3).     X  about 

1600. 
Figs.  281  to  284.     Spores  of  Myxidium  anguiUae.    After  Ishii  (1915,  Fig.  3a).     X1450, 
Figs.  285  and  286.    Myxidium  sp.    After  Mavor  (1915). 
Fig.  285.    A  spore  treated  with  ammonia  water  (1915,  Fig.  3a).     X660. 
Fig.  286.    A  spore  (1915,  Fig.  3b).     X1320. 

Figs.  287  to  290.     Spores  of  Myxidium  gadi.    After  Georg6vitch  (1916,  Figs.  1,  4,  3  and  2). 
Fig.  228.     A  spore  from  Solea  vulgaris. 
Figs.  289  and  290.     Young  spores. 


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Fig.  291.    A  spore  of  Myxidium  glutinosum.    After  Davis  (1917,  Fig.  124).     X1400. 

Figs.  292  and  293.    Spores  of  Myxidium  phyllium.    After  Davis  (1917,  Figs.  126  and  127). 

X2000. 
Figs.  294  and  295.    Spores  of  Myxidium  kagayatnai.    After  Kudo  (1916,  Fig.  2).     X1750 

and  X 1000  respectively. 
Figs.  296  to  307.    Sphaeromyxa  balbianii. 
Fig.  296.    A  trophozoite.    After  Thdohan  (1895,  Fig.  57).     X3. 
Fig.  297.    A  trophozoite.    After  Davis  (1917,  Fig.  128).     X640. 
Fig.  298.    A  trophozoite  in  plasmotomy.    After  Georg6vitch  (1916,  Fig.  15). 
Figs.  299  and  300.    Spores.    After  Th61ohan  (1895,  Figs.  58  and  59).     X1500. 
Fig.  301.    An  end  of  a  spore.    After  Thflohan  (1895,  Fig.  60). 
Fig.  302.    A  spore  treated  with  nitric  acid.    After  Thflohan  (1895,  Fig.  61). 
Fig.  303.    A  spore.    After  Parisi  (1912,  Fig.  4).     X  about  1750. 
Fig.  304.    A  spore.    After  Davis  (1917,  Fig.  130).     X2100. 
Figs.  305  to  307.    Mature  and  young  spores  (Figs.  306  and  307).     After  Georg6vitch  (1916, 

Figs.  17,  20  and  19). 
Figs.  308  to  311.    Sphaeromyxa  immersa.    After  Lutz  (1889:  301). 
Fig.  308.    An  infected  gall-bladder  of  Bufo  aqua  (1889,  Fig.  1).     XL 
Fig,  309.    Spores  (1889,  Figs.  4,  5  and  6). 
Fig.  310.    A  spore  (1889,  Fig.  10).     X600. 
Fig.  311.    A  spore  with  extruded  polar  filaments  (1889,  Fig.  7). 
Figs.  312  to  314.    Spores  of  Sphaeromyxa  incurvata.    After  Doflein  (1898,  Fig.  49).     X 

about  1000. 
Figs.  315  and  316.    Sphaeromyxa  sabrazesi.    After  Schroder  (1907). 
Fig.  315.    A  trophozoite  (1907,  Fig.  1).     X15. 
Fig.  316.    A  cross  section  thru  a  trophozoite  (1907,  Fig.  3).     XlSOO. 


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238  JLUNOIS  BIOLOGICAL  MONOGRAPHS  [476 


EXPLANATION  OF  PLATE 

Figs.  317  to  322.    Spores  of  Sphaeromyxa  sabrazesi. 

Figs.  317  to  319.    After  Laveran  and  Mesnil  (1900,  Figs.  1,  3  and  4).     XlSOO. 

Fig.  318.    A  spore  treated  with  nitric  acid  (1900,  Fig.  3). 

Figs.  320  and  321.    Spores.    After  Schroder  (1907,  Figs.  39  and  41).     X1500. 

Fig.  322.    A  polar  capsule.    After  Schroder  (1907,  Fig.  45). 

Figs.  323  and  324.    Spores  of  Sphaeromyxa  hellandi.    After  Auerbach  (1909,  Fig.  5).     X 

about  1500. 
Figs.  325  and  326.    Spores  of  Sphaeromyxa  exneri.    After  Awerinzew  (1913a,  Fig.  3).     X 

about  365. 
Fig.  327.    A  spore  of  Sphaeromyxa  gasterostei.    After  Georg6vitch  (1916,  Fig.  22). 
Figs.  328  to  331.    Zschokkella  hildae.    After  Auerbach  (1910a  and  1912). 
Fig.  328.    A  monosporous  trophozoite  (1912,  PL  5,  Fig.  2). 
Figs.  329  to  331.    Spores  (1910a,  Fig.  62). 
Figs.  332  and  333.    Spores  of  Zschokkella  ruma.    After  Klokacewa  (1914,  Fig.  2).     X  about 

2500. 
Figs.  334  to  338.    Spores  of  Zschokkella  acheilognathi.    After  Kudo  (1916). 
Figs.  334  to  336.    Different  views  of  fresh  spores  (1916,  Figs.  3d,  3e  and  3f).     X2250. 
Fig.  337.    A  young  spore.    Original.     X2785. 
Fig.  338.    A  stained  spore  (1916,  Fig.  3h).     X2800. 
Figs.  339  and  340.    Spores  of  Zschokkella  globulosa.    After  Davis  (1917,  Figs.  135  and  134). 

X1500. 
Figs.  341  to  343.    Spores  of  Myxosoma  dujardini.    After  Th6Iohan  (1895,  Figs.  90,  91,  and 

89).     X1500. 
Figs.  344  to  347.    Spores  of  Myxosoma  funduli.    After  Kudo  (1918a,  Fig.  A).     X1500. 


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210  ILUNOJS  BIOLOGICAL  MONOGRAPHS  [478 


EXPLANATION  OF  PLATE 

Fig.  348.    A  spore  of  Myxosomai?)  lobatum.    After  Nemeczek  (1911,  Fig.  18).     XlOOO. 

Fig.  349  to  354.    Lentospora  cerebralis.    After  Plehn  (1905).    X1200. 

Fig.  349.     Various  young  ameboid  forms;  stained  (1905,  Textfig.  5), 

Fig.  350.    A  stained  larger  form  (1905,  Textfig.  5). 

Fig.  351.    A  trophozoite  with  two  spores  (1905,  Textfig.  4). 

Fig.  352.    Various  spores  (1905,  Textfig.  2). 

Fig.  353.    A  stained  spore  (1905,  Textfig.  3). 

Fig.  354.    A  spore  with  extruded  polar  filaments  (1905,  Textfig.  2). 

Fig.  355.    A  spore  oi  Lentospora  multiplicata.    After  Reuss  (1906,  Fig.  8).     X1500. 

Figs.  356  to  359.    Spores  of  Lentospora  asymmetrica.    After  Parisi  (1912,  Fig.  7).     X  about 

1500. 
Figs.  360  to  362.    Spores  of  Lentospora  acuta.    After  Fujita  (1912,  Fig,  2).     X2000. 
Figs.  363  and  364.    Spores  of  Myxobolus  piriformis.    After  Thdohan  (1895,  Figs.  117  and 

116).     X1500. 
Figs.  365  and  366.    Spores  of  Myxobolus  unicapsulaius.    After  Miiller  (1841,  Fig.  5). 
Fig.  367.    A  spore  of  Myxobolus  fukrmanni.    After  Auerbach  (1909,  Fig.  lb).     X1840. 
Fig.  368.    A  spore  of  Myxobolus  oculi4eucisci.    After  Trojan  (1909,  Fig.  3).     X2000, 
Figs.  369  and  370.    Spores  of  Myxobolus  toyatnai.    After  Kudo  (Original  and  1917,  Fig.  40). 

X25O0. 
Figs.  371  and  372.    Spores  of  Myxobolus  notatus.    After  Mavor  (1916,  Figs.  6C  and  6B). 

X2600. 
Figs.  373  and  374.    Spores  of  Myxobolus  rohitae.    After  Southwell  and  Prashad  (1918, 

Figs.  26  and  27).     X  about  1720  and  700  respectively. 
Figs.  375  and  376.     Spores  of  Myxobolus  sent.    After  Southwell  and  Prashad  (1918,  Figs. 

29  and  30).     X  about  1700. 
Figs.  377  and  378.    Sports  oi  Myxobolus  misgurni.    Original.     X1500. 


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PLATE  XVI 


242  ILLINOIS  BIOLOGICAL  MONOGRAPHS  i^» 


EXPLANATION  OF  PLATES 

Figs.  379  to  385.    Myxobolus  pfeifferi. 

Figs.  379  and  380.    Parts  of  section  thru  cyst.     After  Keysselitz  (1908a,  PI.  15,  Figs.  1  and  2) . 

Fig.  381.    A  spore.    After  TMlohan  (1895,  Fig.  77).     X1500. 

Fig.  382.    An  optical  section  of  spore.    After  Keysselitz  (1908a,  Fig.  A). 

Fig.  383.    A  spore  treated  with  Lugol's  solution.     After  Keysselitz  (1908a,  PI.  14:  Fig.  92). 

Fig.  384.    A  spore  with  extruded  filaments.    After  Keysselitz  (1908a,  Fig.  C). 

Fig.  385.    A  stained  young  spore.    After  Keysselitz  (1908a,  PI.  14,  Fig.  81). 

Fig.  386.    A  spore  of  Myxobolus  dispar.    After  Thdlohan  (1895,  Fig.  86).     X  about  1500. 

Figs.  387  to  389.     Spores  of  Myxobolus  ellipsoides.     Fig.  389.    An  abnormal  spore  with  six 

polar  capsules.    After  Thelohan  (1895,  Figs.  112,  113  and  115). 
Fig.  390.    A  part  of  the  infected  intestine  of  Mugil  auratus,  showing  cysts  of  Myxobolus 

exiguus.    After  Parisi  (1912,  Fig.  9e).     X  about  3. 
Fig.  391.    A  spore  of  Myxobolus  exiguus.    After  Thelohan  (1895,  Fig.  98).     X  1500. 
Figs.  392  to  395.     Spores  of  Myxobolus  exiguus.    After  Parisi  (1912,  Fig.  9).     X2500. 
Fig.  396.    A  spore  of  Myxobolus  oviformis.    After  Th61ohan  (1895,  Fig.  81).     X1500. 
Figs.  397  to  398.     Spores  of  Myxobolus  miiUeri.    After  Thdohan  (1895,  Figs.  96  and  97). 

X1500. 
Figs.  399  and  400.     Spores  of  Myxobolus  millleri.    After  Biitschli  (1881,  Figs.  1  and  2). 
Fig.  401.    A  spore  of  Myxobolus  millleri  in  cone,  sulphuric  acid.    After  Biitschli  (1881,  Fig.  6). 
Figs.  402  and  403.    Abnormal  spores  of  Myxobolus  millleri.    After  Stitschli  (1881,  Figs.  9 

and  8). 
Figs.  404  and  405.     Spores  of  Myxobolus  lintoni.    After  Linton  (1891,  Figs.  3  and  5). 
Fig.  406.    Diagram  of  the  cross  section  of  a  spore  of  Myxobolus  lintoni.    After  Linton  (1891, 

Fig.  8). 
Fig.  407.    A  spore  of  Myxobolus  linloni  with  extruded  polar  filaments.    After  Linton  (1891, 

Fig.  10). 
Fig.  408.    A  stained  spore  of  Myxobolus  lintoni.    After  Gurley  (1894,  PI.  26,  Fig.  7).     X 

about  2000.  • 

Figs.  409  and  410.     Spores  of  Myxobolus  globosus.    After  Gurley  (1894,  PI.  26,  Fig.  7). 

X  about  2900. 
Fig.  411.    Sp>ores  of  Myxobolus  inaequalis.    After  Miiller  (1841,  Fig.  6). 


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PLATE  XVII 


2M  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [482 


EXPLANATION  OF  PLATE 

Figs.  412  to  416.  Spores  of  Myxobolus  oblongus.  Figs.  412  to  414.  After  Gurley  (1894, 
PI.  26,  Fig.  6).     X2300.    Figs.  415  and  416.    After  MiiUer  (1841,  Fig.  9). 

Figs.  417  and  418.    Spores  of  Myxobolus  transovdis.    After  Gurley  (1894,  PI.  29,  Fig.  1). 

Figs.  419  and  420.    Spores  of  Myxobolus  obesus.    After  Balbiani  (1884,  Fig.  39). 

Fig.  421.    Spores  of  Myxobolus  cycloides.    After  Miiller  (1841,  Fig.  3). 

Figs.  422  and  423.    Spor^  oi  Myxobolus  anurus.    After  Cohn  (1895,  Fig.  25).     X1500. 

Fig.  424.    Spores  of  Myxobolus  sp.  X700.    After  Butschli  (1882,  PL  36,  Fig.  23). 

Fig.  425.    Myxobolus  sp.    After  Gurley  (1894,  PL  28:  Fig.  4a).     X  about  1500. 

Figs.  426  to  429.    Spores  of  Myxobolus  sp.    After  Miiller  (1841,  Fig.  4). 

Fig.  430.    A  vegetative  form  of  Myxobolus  cyprini.    After  Doflein  (1898,  Fig.  112). 

Figs.  431  and  432.    Spores  of  Myxobolus  cyprini.    After  Doflein  (1898,  Figs.  113  to  115). 

Figs.  433  to  436.    Myxobolus  neurobius.    After  Schuberg  and  Schroder  (1905). 

Figs.  433  and  434.  Longitudinal  and  transverse  sections  thru  infected  nerve  fibres  (1905, 
Figs.  2  and  4a).     X520. 

Figs.  435  and  436.    Sp>ores  (1905,  Figs.  5  and  6).    Comp.  oc.  12  and  imm.  obj.  2mm. 

Figs.  437  to  441.    Myxobolus  aeglefini.    After  Auerbach  (1906a). 

Fig.  437.    A  cyst  in  the  sclerotic  cartilage  of  the  eye  of  Gadus  aeglefini  (1906a,  Fig.  2). 

Figs.  438  and  439.    Spores  (1906a,  Figs.  5a  and  3d).     Xabout  1320. 

Figs.  440  and  441.    Abnormal  spores  (1906a,  Figs.  5b  and  5c).     Xabout  1320. 

Figs.  442  to  445.    Myxobolus  gigas.    After  Auerbach  (1906b). 

Fig.  442.    A  part  of  the  section  of  a  cyst  (1906b,  Fig.  1). 

Figs.  443  to  445.    Spores  (1906b,  Figs.  3a,  3c,  5  and  3b).     X  about  850. 

Figs.  446  and  447.    Spores  of  Myxobolus  volgensis.    After  Reuss  (1906,  Fig.  1).     X2000. 


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4831  STUDIES  ON  MYXOSFORIDIA—KUDO  3^ 


PLATE  XVIII 


246  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [484 


EXPLANATION  OF  PLATE 

Fig.  448.    The  branchia  of  Lucioperca  volgensis  with  cysts  of  Myxobolus  volgensis.    After 

Reuss  (1906,  Fig.  2).     X2.25. 
Fig.  449.    A  spore  of  Myxobolus  scardinii.    After  Reuss  (1906,  Fig.  3).     X1500. 
Fig.  450.    Air  bladder  of  Scardinius  erythrophtkalmus  with  the  cysts  of  Myxobolus  pkyso- 

philus.    After  Reuss  (1906,  Fig.  5).     X2. 
Fig.  451.    A  spore  of  Myxobolus  physophilus.    After  Reuss  (1906,  Fig.  4).     X1500. 
Fig.  452.    A  sport  of  Myxobolus  macrocapsular is.    After  Reuss  (1906,  Fig.  6).     X1500. 
Fig.  453.    A  spore  of  Myxobolus  sandrae.    After  Reuss  (1906,  Fig.  7).     X2000. 
Fig.  454.    A  spoK  ol  Myxobolus  bramae.    After  Reuss  (1906,  Fig.  9).     X1500. 
Fig.  455.    A  spore  ot  Myxobolus  balleri.    After  Reuss  (1906,  Fig.  10).     X1500. 
Fig.  456.    A  spore  of  Myxobolus  cyprinicola.    After  Reuss  (1906,  Fig.  11).     X1500. 
Figs.  457-459.    Myxobolus  squamae.    After  Keysselitz  (1908a). 
Fig.  457.    A  part  of  the  infected  scale  (1908a,  Fig.  G.) 
Fig.  458.    A  spore  treated  with  Lugol's  solution  (1908a,  PI.  14,  Fig.  94). 
Fig.  459.    A  stained  spore  (1908a,  PL  14,  Fig.  96). 

Figs,  460  and  461'.    Spores  of  Myxobolus  cordis.    After  Keysselitz  (1908a). 
Fig.  460.    A  spore  treated  with  Lugol's  solution  (1908a,  PI.  16,  Fig.  16). 
Fig.  461.    A  strained  spore  (1908a,  Fig.  B  on  page  281). 
Figs.  462  to  464.     Spores  of  Myxobolus  tnusculi.    After  Keysselitz  (1908a). 
Fig.  462.    A  spore  treated  with  Lugol's  solution  (1908a,  PI.  15,  Fig,  13). 
Fig.  463  and  464.    Stained  spores  (1908a,  Figs.  D  and  E  on  page  286). 
Fig.  465.    Sports  oi  Myxobolus  sp.    After  Wegener  (1910,  Fig.  44).     X1050. 
Fig.  466.    A  sport  oi  Myxobolus  per magnus.    After  Wegener  (1910,  Fig.  45).     X1050. 
Fig.  467.    Spores  of  Myxobolus  rotundus.    After  Nemeczek  (1911,  Figs.  10  and  11).     XlOOO. 
Fig.  468.    Spores  of  Myxobolus  minutus.    After  Nemeczek  (1911,  Figs.  16  and  17).     XlOOO. 
Figs.  469  and  470.    Spores  of  Myxobolus  magnus.    After  Awerinzew  (1913,  76).     X  about 

340. 
Figs.  471  to  473.    Spores  of  Myxobolus  carassii.    After  Klokacewa  (1914,  Fig.  1).     X  about 

2400. 


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PLATE  XIX 


248  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [186 


EXPLANATION  OF  PLATE 

Figs.  474  to  476,    Myxobolus  funduli.    After  Hahn  (1915  and  1917). 

Fig.  474.    A  cyst  from  the  gill  filament  (1917,  Fig.  1). 

Fig.  475.    A  stained  spore  (1915,  Fig.  28).     X2000. 

Fig.  476.    Diagram  of  the  cross  section  of  a  spore  (1915,  Fig.  30). 

Fig.  477.    A  spore  of  Myxobolus  pleuronectidae.    After  Hahn  (1917a,  Fig.  2).     X157S. 

Fig.  478.    A  spore  of  Myxobolus  capsulatus.    After  Davis  (1917,  Fig.  139).     X1500. 

Figs.  479  and  480.  Spores  of  Myxobolus  nodularis.  After  Southwell  and  Prashad  (1918, 
PI.  11,  Figs.  34  and  35).     X  about  1450. 

Fig.  481.    A  sf)ore  of  Myxobolus  miyairii.    After  Miyairi  (1909,  Fig.  14). 

Figs.  482  to  485.     Spores  of  Myxobolus  koi.    Original.     X 1300. 

Figs.  482  to  484.    Different  views. 

Fig.  485.    A  spore  stained  with  Giemsa's  mixture. 

Figs.  486  and  487.    Henneguya  psorospermica.    After  Th€lohan  (1895). 

Fig.  486.    A  cross  section  thru  branchial  lamella  of  Esox  lucius  with  a  cyst  (1895,  Fig.  82). 

Fig.  487.    Two  spores  (1895,  Figs.  83  and  84).     X  about  1000. 

Figs.  488  and  489.    Henneguya  minuta.    After  Cohn  (1895). 

Fig.  488.    A  longitudinal  section  of  an  infected  branchial  lamella  (1895,  Fig.  29.) 

Fig.  489.    Two  spores.    One  with  two  vacuoIes(?)  (1895,  Fig.  30).     X  about  450. 

Fig.  490  and  491.    Spores  of  Henneguya  oviperda.    After  Cohn  (1895,  Fig.  31). 

Fig.  491.    A  spore  with  extruded  "starren  Faden"  and  polar  filaments. 

Figs.  492  and  493.    Henneguya  lobosa.    After  Cohn. 

Fig.  492.    An  external  view  of  the  parasite  on  the  gill  (1895,  Fig.  18). 

Fig.  493.    Two  spores  and  one  unseparated  young  spores  (1895,  Fig.  21). 

Figs.  494  and  495.    Henneguya  media.    After  TMlohan  (1890b). 

Fig.  494.    A  sporoblast  in  the  ovary  of  Gasterosteus,  with  one  spore  (1890b,  Fig.  18). 

Fig.  495.    Spores  (1890b,  Fig.  1). 

Fig.  496.  The  peripheral  portion  of  a  section  of  a  cyst  of  Henneguya  psorospermica,  show- 
ing the  characteristic  structure.    After  Th^lohan  (1895 :  237). 

Figs.  497  to  499.    Spores  of  Henneguya  schizura.    After  Muller  (1841,  Fig.  1). 

Figs.  500  to  503.    Spores  of  Henneguya  creplini.    After  Creplin  (1842,  Figs.  B,  E,  A  and  C). 

Fig.  504.    Spores  of  Henneguya  linearis.    After  Miiller  (1841,  Fig.  10). 


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250  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [488 


EXPLANATION  OF  PLATE 

Fig.  505.    Spores  of  Henneguya  Gurleyi.    After  Gurley  (1894,  PI.  ii,  Figs.  8c,  6  and  7). 

X  about  3100. 
Fig.  506.    Spores  of  Henneguya  strongylura.    After  Muller  (1841,  Fig.  2). 
Fig.  507.    Spores  of  Henneguya  tnonura.    After  Ryder  (1880,  Figs,  Ic  and  2d). 
Fig.  508.    Sp>ores  of  Henneguya  kolesnikovi.    After  Kolesnikov  from  Gurley  (1894,  PL  35, 

Fig.  7). 
Figs.  509  to  512,    Henneguya  macrura.    After  Gurley  (1894).     X  about  2100. 
Figs.  509  and  510.    Spores  (1894,  PI.  32,  Fig.  5,  PI.  33,  Fig.  1). 

Fig.  511.    A  spore  treated  with  iodine,  showing  the  "beading  of  the  tail"  (1894,  PI.  33,  Fig.  3). 
Fig.  512.    A  tail  separated  from  the  main  part  by  iodine  (1894,  PI.  ii,  Fig.  4). 
Fig.  513.    Spores  of  Henneguya  zschokkei.    After  Zschokke  (1898,  Figs.  2  and  1). 
Fig.  514.    Spores  of  Henneguya  sp.    After  Benecke  from  Gurley  (1894,  PI.  29,  Fig.  8). 
Fig.  515.    Spore  of  Henneguya  tenuis.    After  Vaney  and  Conte  (1901,  Fig.  2). 
Figs.  516  and  517.    Spores  of  Henneguya  nusslini.    After  Schuberg  and  SchrSder  (1905, 

Figs.  13  and  14),    Comp.  oc.  12  and  obj.  2mm. 
Figs.  518  to  523.    Henneguya  legeri.    After  C6pMe  (1913). 
Figs.  518  to  521.    Trophozoites  (1913,  Figs.  2,  23,  15  and  25).     X900. 
Fig.  519.    A  trophozoite  in  division  (1913,  Fig.  23),     X900. 
Fig.  521.    A  trophozoite  stained  with  iron  hematoxylin  (1913,  Fig.  25).     X900. 
Fig.  522.    An  elongated  spore  (1913,  Fig.  26).     X900, 
Fig,  523,    An  ovoidal  spore  (1913,  Fig.  24),     X450. 
Fig.  524.    A  spore  of  Henneguya  miyairii.    After  Miyairi  (1909,  Fig,  11). 


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STUDIES  OW  MTZOSPOEIDI A— KUDO  2H 


PLATE  XXI 


252  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [490 


EXPLANATION  OF  PLATE 

Figs.  525  and  526.    Spores  of  Eenneguya  acerinae.    After  SchrSder  (1906,  Figs.  5  and  6). 

X1650. 
Fi^.  527  to  535.    Eenneguya  giganUa.    After  Nemeczek  (1911).     XlOOO. 
Fig.  527.    A  mature  spore  with  extruded  polar  filaments  (1911,  Fig.  1). 
Figs.  528  to  535.    Stages  in  development  of  spores  (1911,  Figs.  2  to  9). 
Figs.  536  to  539.    Spores  of  Henneguya(?)  sp.    After  Nemeczek  (1911,  Figs.  12  to  15). 

XlOOO. 
Figs.  540  to  543.    Eenneguya  gasterostet.    After  Parisi  (1912).     X  about  1500. 
Fig.  540.    A  disporous  trophozoite  (1912,  Fig.  lOd). 
Figs.  541  and  542.    Two  spores  (1912,  Figs.  lOf  and  lOe). 
Fig.  543.    A  young  sp>ore  (1912,  Fig.  10c). 
Figs.  544  and  545.    Spores  of  Eenneguya  neapolilana.    After  Parisi  (1912,  Fig.  11).     X 

about  1500. 
Figs.  546  to  549.    Various  trophozoites  of  Eenneguya  mictospora.    OriginaL 
Fig.  546.    A  monosporous  trophozoite  with  a  yoimg  spore.     X950. 
Figs.  547  and  549.    Trophozoites  in  vivo.     X650. 
Fig.  548.    A  stained  binucleated  yoimg  trophozoite.     X1700. 


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254  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [492 


EXPLANATION  OF  PLATE 

Figs.  550  to  557.    Henneguya  mictospora.    Original. 

Fig.  550.    A  stained  trophozoite.     X1700. 

Figs.  551  to  553.    Three  different  stages  of  development  of  disporous  trophozoites.    Giemsa. 

X1700. 
Figs.  554  and  555.    Two  stages  of  monosporous  trophozoites.    Giemsa.     X1700. 
Figs.  556  and  557.    Different  views  of  mature  spores  in  vivo.     X2000. 
Figs.  558  and  559.    Henneguya  wisconsinensis.    After  Mavor  and  Strasser  (1916). 
Fig,  558.    A  trophozoite  in  vivo  (1916,  Fig.  la).     X570. 
Fig.  559.    A  fresh  spore  (1916,  Fig.  3d).     X4000. 

Figs  560  and  561.    Trophozoites  oi  Chloromyieum  catostomi.    Original.     X1500. 
Figs.  562  to  565.    Chlorotnyxum  clupeidae.    Original  drawn  from  Dr.  Tyzzer's  smears  which 

were  restained.     X2360. 
Fig.  562.    Anterior  end  view  of  two  spores  in  preserved  and  decolorized  smears. 
Fig.  563.    The  same  views  of  three  spores  restained  with  Giemsa  mixture. 
Fig.  564.    Front  view  of  a  preserved  and  decolorized  spore. 
Fig.  565.    The  same  views  of  two  spores  restained  with  Giemsa's  mixture. 
Figs.  566  to  572.    Spores  of  Myxobolus  orbiculatus.    Original. 

Figs.  566,  569  and  570.    Different  views  of  normal  spores  in  preserved  specimen.     X 1500. 
Figs.  567  and  568.    Abnormal  spores.     X1500. 
Fig.  571.    A  spore  stained  with  Lugol's  solution.     X1500. 
Kg.  572.    A  spore  stained  with  Giemsa's  mixture.     X2360. 


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2M  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [494 


EXPLANATION  OF  PLATE 

Figs.  573  to  576.    Myxobolus  orbiculatus.    Original 

Fig.  573.    Young  vegetative  forms  in  the  muscle  fibres.     From  a  section  stained  with  Heid- 

enhain's  iron  hematoxylin.     X900. 
Figs.  574  and  575.    Two  vegetative  forms  under  higher  magnifications.     X1500. 
Fig.  576.    A  cross  section  thru  the  infected  muscle  fibres,  stained  with  Heidenhain's  iron 

hematoxylin.     X900. 
Figs.  577  to  581.    Hoferellus  cyprini.    After  Doflein  (1898). 
Figs.  577  and  578.    Two  vegetative  forms  (1898,  Figs.  106  and  105). 
Figs.  579  to  581.    Spores  (1898,  Figs.  108  and  107). 
Figs.  582  and  583.    Genus  incert.    merlucii.    After  Perugia  from  Gurley  (1894,  PI.  29, 

Figs.  2  and  7). 
Figs.  584  and  585.    Genus  incert.  congri.    After  Perugia  from  Gurley  (1894,  PL  6,  Figs.  7 

and  3). 
Figs.  586  and  587.    Genus  et  species  incertae.    After  Mavor  (1915). 
Fig.  586.    A  trophozoite  with  attached  Ceratomyxa  acadiensis  (1915,  Fig.  8).     X830. 
Fig.  587.    A  trophozoite  (1915,  Fig.  6). 
Figs.  588  and  589.    Trophozoites  of  genus  et  species  incertae.    After  Mavor  (1916a,  Figs. 

3d  and  3b).     X660. 
Fig.  586.    A  trophozoite  with  attached  Ceratomyxa  acadiensis  (1915,  Fig.  8).     X830. 
Fig.  587.    A  trophozoite  (1915,  Fig.  6). 
Figs.  588  and  589.    Trophozoites  of  genus  et  species  incertae.    After  Mavor  (1916a,  Figs. 

3d  and  3b).     X660. 
Fig.  590.     SjK)res  of  genus  et  species  incertae.    After  Linton  (1891a,  Fig.  2). 
Figs.  591  to  593.    Myocobolus  hylae.    After  Johnston  and  Bancroft  (1918). 
Fig.  591.    A  transverse  section  of  a  heavily  infected  testis  of  Hyla  aurea,  (1918,  Rg.  1). 

X  about  11. 
Fig.  592.    Different  views  of  normal  spores,  stained  (1918,  Fig.  3).     X  about  800. 
Fig.  593.    Abnormal  spores  (1918,  Fig.  4).     X  about  800. 
Figs.  594  to  596.    Lentospora  dermaiobia.    After  Ishii  (1915). 
Fig.  594.    A  part  of  the  infected  skin  of  the  host  (1915,  Fig.  2).     X140. 
Figs.  595  and  596.    Different  views  of  spore  (1915,  Figs.  4  and  3).     X1450. 
Figs.  597  to  601.    Myxobolus  discrepans.    Original. 
Fig.  597.    An  infected  branchial  lamella  showing  the  cysts  of  various  size  and  form.    X 

about  4. 
Figs.  598  to  601.    Unstained  preserved  spores,  showing  different  views.     X  about  1500. 


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258  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [496 


EXPLANATION  OF  PLATE 

Figs.  602  to  621.  MUraspora  elongate.  Original.  X 1500,  except  Figs.  620  and  621,  X2500. 
Giemsa  staining,  unless  otherwise  stated. 

Fig.  602.  A  trophozoite  showing  various  nuclei  and  sporoblasts  at  different  stages  of  devel- 
opment. 

Fig.  603.    A  trophozoite  with  a  mature  and  a  young  spore.    Iron  hematoxylin. 

Fig.  604.    A  trophozoite  with  two  mature  spores. 

Figs.  605  to  609.     Formation  and  development  of  sporoblasts. 

Figs.  610  to  613.     Developing  spores.     Fig.  612;  Delafield's  hematoxylin. 

Fig.  614.    A  surface  view  of  a  preserved  spore. 

Fig.  615.    An  optical  section  of  a  preserved  spore. 

Fig.  616.     Front  view  of  a  stained  spore. 

Fig.  617.    Lateral  view  of  a  spore  stained  with  Heidenhain's  iron  hematoxylin. 

Fig.  618.    A  slightly  elongated  spore. 

Fig.  619.    An  abnormal  spore. 

Fig.  620.    A  longitudinal  section  thru  a  polar  capsule. 

Fig.  621.    An  oblique  view  of  a  polar  capsule,  showing  the  spirally  coiled  polar  filament. 

Figs.  622  to  627.    Myxidium  americanum.    Original.     X1500. 

Figs.  622  and  623.    Two  young  trophozoites.     Giemsa. 

Fig.  624.    A  sporulating  trophozoite  in  imstained  preserved  state. 

Fig.  625.    A  spore  in  preserved  state. 

Fig.  626.    A  fresh  spore  treated  with  potassium  hydrate  solution. 

Fig.  627.    A  spore  stained  with  Giemsa's  mixture. 

Figs.  628  to  631.    Myxobolus  mesentericus.    Original.     X1500. 

Figs.  628  and  629.     Front  and  lateral  views  of  unstained  preserved  spores. 

Fig.  630.    A  Giemsa  stained  spore. 

Fig.  631.  A  spore  with  extruded  polar  filament  (potassium  hydrate),  stained  with  Giemsa's 
mixture. 


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200  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [498 


EXPLANATION  OF  PLATE 

Figs.  632  to  642.    CUoromyxum  wardi.    OriginaL 

Fig.  632.    A  3roung  trophozoite.    Smear  and  Giemsa,     XlSOO. 

Fig.  633.    A  sporulating  trophozoite.    Giemsa.     X 1500. 

Figs.  634  to  637.    Surface  views  and  optical  sections  of  four  unstained  spores,  showing  the 

sutural  ridge  and  the  fine  striations  on  the  shelL     X 1500. 
Fig.  638.    A  polar  view  of  unstained  spore,     X2360. 
Figs.  640.    Surface  view  and  optical  section  of  a  single  spore.     X2360. 
Fig.  641.    A  front  view  of  an  unstained  spore.     X2360. 
Fig.  642.    A  Giemsa  stained  spore.     X1500. 
Figs.  643  to  649.    Myxoholus  aureatus.    After  Ward  (1919). 
Fig.  643.    A  fresh  spore  (1919,  Fig.  Aa). 

Figs.  644  and  645.    Fresh  spores  kept  for  24  hours  or  more  in  water  (1919,  Fig.  Aa). 
Fig.  646.    A  preserved  unstained  spore  (1919,  Fig.  Ba). 

Fig.  647.    A  spore  stained  with  Delafield's  hematoxylin  (1919,  Fig.  Bd).     X1500. 
Fig.  648.    A  spore  with  an  extruded  polar  filament  from  section  preparation  stained  with 

Giemsa's  mixture  (1919,  Fig.  Bg).     X1500. 
Fig.  649.    A  yoimg  spore,  stained  with  Giemsa's  mixture  (1919,  Fig.  Bi).     X1500. 
Figs.  650  to  653.    Henneguya  brachyura.    After  Ward  (1919).     X1500. 
Fig.  650.    A  young  spore  stained  with  Giemsa's  mixture  (1919,  Fig.  Ce). 
Figs.  651  and  652.    Front  and  lateral  views  of  spore  (1919:  F^.  Cb  and  Cc). 
Fig.  653.    A  detached  taU  (1919,  Fig.  Cf). 

Figs.  654  to  656.  Henneguya  salminicola.  After  Ward  (1919).  X1500. 
Fig.  654.  A  young  spore  stained  with  Giemsa's  mixture  (1919,  Fig.  Fc). 
Figs.  655  and  656.    Front  and  lateral  views  of  unstained  preserved  spores. 


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INDEX 


African  Myxosporidia,  14 

Air  bladder,  Mj^osporidia  parasitic  in,  39 

Alaskan  Myxosporidia,  16 

Alimentary  canal,  Myxosporidia  parasitic  in, 

39,41 
Amphibian  hosts  of  Myxosporidia,  36 
Annelida  as  host  of  Mjrxosporidian,  25 
Anterior  end  of  spore,  47,  50 

process  of  spore,  47,  50,  51 
Asiatic  Myxosporidia,  13 
Australian  Myxosporidian,  14 
Austrian  Myxosporidia,  24 
Blood  vessel,  Mj^tosporidia  parasitic  in,  39 
Body  cavity,  Myxosporidia  parasitic  in,  37, 

43,45 
Bone,  Myxosporidia  parasitic  in,  39 
Branchia,  Myxosporidia  parasitic  in,  38 
Breadth  of  spore,  50 
Burma,  Myxosporidia  of,  14 
Canadian  M5Tcosporidia,  16 
Capsulogenous  cell  of  spore,  50 
Cartilage,  Myxosporidia  parasitic  in,  39 
Ceratomyxa,  9,  56,  65,  178,  180 
Ceratomyxa  abbreviata,  76 

acadiensis,  71 

agglomerata,  77 

aggregata,  79 

amorpha,  78 

appendiculata,  67 

arcuata,  65 

attenuaia,  75 

coris,  72 

drepanopsettae,  70 

flagellifera,  77 

globurifera,  67 

herouardl,  72 

inaequdis,  68 

Unospora,  69 

lunata,  76 

mesospora,  73 

tnonospora,  78 

navicularia,  80 

pallida,  67 

ramosa,  69 

recurvata,  75 

reticularis,  68 

spari,  70 

sp.  (?)  Awerinzew,  71 


sp.  (?)  Awerinzew,  71 
sp.  Georg^vitch,  72 
sphaerulosa,  66 
sphairophora,  73 
spinosa,  80 
streptospora,  79 
taenia,  74 
truncata,  67 
tylosuri,  ^0 
undtdata,  79 
Ceratom)^dae,  9,  56,  60,  178 
Chloromyxidae,  10,  57,  87,  178 
Chloromyxum,  10,  57,  87,  178,  183 
CMoromyxum  catostomi,  98 
cattdatutn,  88 
dupeidae,  94 
cristatum,  91 
diploxys,  90 
dubium,  91 
fluviatile,  89 
fujitai,  93 
funduli,  93 
granulosutn,  96 
yfeoi,  92 
leydigi,  87 
magnum,  92 
misgurni,  93 
tnucronaium,  89 
^ro<e«,  90 
quadratum,  88 
sp.  Awerinzew,  91 
thymalli,  92 
trijugum,  96 
truttae,  90 
wardi,  99 
Classification  of  Myxosporidia  after 
Auerbach,  52 
Davis,  54 
Doflein,  52 
Kudo,  56 
Parisi,  54 
Poche,  54 
Thelohan,  52 
Connettive  tissue,  Myxosporidia  parasitic  in, 

37 
Cyst,  50 
Disporea,  52,  54 
Disporous,  50 


262 


ILUNOIS  BIOLOGICAL  MONOGRAPHS 


[500 


England,  Myxosporidia  of,  22 
European,  M)aosporidia,  17 
Eurysporea,  9,  56,  60 
Eye,  Myxosporidia  parasitic  in,  38 
Fish  as  hosts  of  Myxosporidia,  25 
fresh  water,  44 
marine,  44 
Foramen  of  polar  capsule,  47,  48,  50 
France,  M3^osporidia  of,  18 
Front  view  of  spore,  47,  50 
Gall  bladder,  Myxosporidia  parasitic  in,  39 
Gemmule,  50 
German  MjTsosporidia,  20 
Heart,  Myxosporidia  parasitic  in,  39 
Henneguya,  11,  59,  158,  179,  193 
Henneguya  acerinae,  167 

brachyura,  171 

brevis,  162 

creplini,  162 

gasterostei,  169 

gigantea,  168 

gurleyi,  163 

kolesnikovi,  164 

legeri,  166 

linearis,  163 

lobosa,  161 

macrura,  164 

media,  161 

mictospora,  173 

minuta,  160 

miyairii,  172 

monura,  164 

neapolitana,  170 

nilsslini,  166 

oviperda,  160 

peri-intestinalis,  161 

psoros  per  mica,  158 

salminicola,  171 

schizura,  162 

sp.  (Gurley)  Labb6, 165 

sp.  (Gurley)  Labb6, 165 

sp.  (?)  Nemeczek,  169 

strongylura,  163 

tenuis,  166 

texta,  159 

wisconsinensis,  170 

zschokkei,  165 
HofereUus,  12,  59,  173,  179 
Hoferettus  cyprini,  174 
Indian  Myxosporidia,  14 
Insect  as  host  of  Myxosporidia,  25 
Int^ument,  Myxosporidia  parasitic  in,  37 


Intercapsular  appendix  of  spore,  47 
Intestine,  Mj^osporidia  parasitic  in,  41 
lodinophilous  vacuole,  47,  50 
Italian  Myxosporidia,  17 
Kamtschatka,  Myxosporidian  of,  14 
Klidney,  Myxosporidia  parasitic  in,  42 
Lateral  process  of  spore,  50 
Length  of  spore,  50 
Lentospora,  11,  58,  125,  179,  189 
Lentospora  acuta,  127 

asymmetrica,  126 

cerebralis,  125 

dermatobia,  127 

encephalina,  126 

multiplicata,  126 
Leptotheca,  9,  56,  60,  178,  179 
Leptotheca  agUis,  60 

dongaia,  60 

fusiformis,  63 

glomerosa,  65 

hepseti,  62 

informis,  63 

lobosa,  64 

longipes,  63 

macrospora,  62 

parva,  61 

perlata,  62 

polymorpha,  61 

renicola,  61 

scissura,  64 

sp.  Awerinzew,  62 
Liver,  Myxosporidia  parasitic  in,  39 
Longitudinal  striation  on  spore,  50 
Mesentery,  Myxosporidia  parasitic  in,  43 
Mesoplasm,  50 
Mictosporea,  52,  54 
Mictosporous,  50 
Mitraspora,  9,  56,  84,  178,  183 
Mitraspora  caudata,  85 
cyprini,  84 
elongata,  85 
Monaco,  Myxosporidia  of,  17 
Monosporous,  50 

Muscle,  Myxosporidia  parasitic  in,  38 
Myxidiidae,  10,  57,  106,  179 
Myxidium,  10,  58,  107,  179,  186 
Myxidium  americanum,  117 

anguiUae,  114 

barbatulae,  110 

bergense,  112 

danilewskyi,  109 

depressum,  114 


•  501] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


263 


gadi,  115 
giardi,  110 
giganteum,  110 
glutinosum,  115 
histophilum,  109 
incurvatum,  108 
infiatum,  111 
kagayamai,  117 
lieberkuhni,  107 
mackiet,  112 
macrocapstdare,  113 
oviforme,  114 
pfeifferi,  111 
phyllium,  116 
sp.  Awerinzew,  113 
sp.  Gurley,  109 
sp.  Mavor,  115 
sphaericum,  109 
striatum,  116 
MyxoboUdae,  11,  58,  128,  179 
Myxobolus,  11,  58,  128,  179,  189 
Myxobolus  aeglefini,  144 
anurus,  142 
aureatus,  154 
6a//er>,  147 
bramae,  147 
capsulatus,  152 
carassii,  ISO 
cordis,  148 
cycloides,  140 
cyprini,  143 
cyprinicola,  147 
discrepans,  156 
dispar,  135 
eUipsoides,  136 
exiguus,  136 
fuhrmanni,  130 
fundtdi,  151 
gtg<w,  145 
globosus,  139 
Ay/oe,  153 
inaequalis,  135 
^o»,  155 
lintoni,  138 
tnacrocapsularis,  146 
magnus,  150 
mesentericus,  157 
minutus,  150 
tnisgumi,  133 
miyairii,  155 
/»tf//eft,  128 
musculi,  148 


neurobins,  144 
nodularis,  153 
notatus,  131 
obesus,  140 
oblongus,  139 
oculi-leucisci,  130 
orbiculatus,  155 
oviformis,  137 
Permagnus,  149 
pfeifferi,  133 
physophilus,  146 
piriformis,  129 
pleuronectidae,  152 
rohitae,  132 
rotundus,  149 
sandrae,  146 
scardinii,  146 
■yewj,  132 
sp.  Gurley,  142 
sp.  Gurley,  142 
sp.  Gurley,  143 
sp.  Kudo,  132 
sp.  Lebzelter,  150 
sp.  Miyairi,  149 
sp.  Southwell,  151 
sp.  Wegener,  149 
sphaeralis,  141 
squamae,  147 
toyamai,  131 
transovalis,  139 
unicapsulatus,  129 
volgensis,  145 
Myxoproteus,  9,  56,  81,  179,  183 
Myxoproteus  ambiguus,  81 
cordiformis,  81 
cornutus,  82 
Myxosoma,  11,  58,  123,  179,  189 
Myxosoma  dujardini,  124 
funduli,  125 
?  lobatum,  124 
Myxosomatidae,  11,  58,  123,  179 
M3^osporidia  in  air  bladder,  39 

alimentary  canal,  39,  41 

Amphibia,  36 

Annelida,  25 

blood  vessel,  39 

body  cavity,  37,  43,  45 

bone,  39 

branchia,  38 

cartilage,  39 

connective  tissue,  37 

eye,  38 


264 


ILUNOIS  BIOLOGICAL  MONOGRAPHS 


[502 


gall  bladder,  39 
heart,  39 
Insecta,  25 
integument,  37 
intestine,  41 
kidney,  41 
liver,  39 
mesentery,  41 
muscle,  38 
nervous  tissue,  39 
organs  of  host,  37 
ovary,  41 
Pisces,  25 
pyloric  coecum,  39 
Reptilia,  36 
spleen,  41 
stomach,  39 
testis,  41 
tissue,  37,  43,  45 
unknown  seat,  41 
urinary  bladder,  41 
of  Africa,  14 
Asia,  13 

Burma,  14 

India,  14 

Japan,  13 

Kamtschatka,  14 

Nippon, 13 
Australia,  14 
Europ>e,  17 

Austria,  24 

England,  22 

France,  18 

Germany,  20 

Italy,  17 

Monaco,  17 

Netherland,  22 

Norway,  22 

Russia,  24 

Serbia,  24 

Switzerland,  23 
North  America,  15 

Alaska,  16 

Canada,  16 

United  States,  15 
South  America,  16 
unknown  genera  and  species, 

12,  174 
Nervous  tissue,  Myxosporidia  parasitic  in,  39 
Netherland,  Myxosporidian  of,  22 
New  species  listed,  196 


Nipjwn,  Myxosporidia  of,  13 

North  American  Myxospwridia,  15 

Norwegian  Myxosporidia,  22 

Organs  of  host  infected  by  Myxosporidia,  list 

of,  37 
Ovary,  Myxosporidia  parasitic  in,  41 
Pansporoblast,  50 
Plasmogamy,  51 
Plasmotomy,  51 
Platysporea,  10,  57,  106,  179 
Polar  capsule,  48,  51 
filament,  48,  51 
Polysporea,  52,  54 
Polysporous,  51 
Posterior  filament,  47,  51 
process,  47,  51 
Pyloric  coecum,  Myxosporidia  parasitic  in,  39 
Reptilian  hosts  of  Myxosporidia,  36 
Ridge  of  spore,  51 
Russian  Myxosp)oridia,  24 
Serbian  Myxosporidian,  24 
Shell  of  spore,  47,  51 

-valve,  47,  51 
Sinuolinea,  10,  57,  104,  178,  186 
Siniwlinea  arborescens,  105 
brachiophora,  106 
capsularis,  105 
dimorpha,  104 
opacita,  106 
South  American  Mj^osporidia,  16 
Species,  scheme  of  description  of,  7 
Sphaeromyxa,  10,  58,  118,  179,  188 
Sphaeromyxa  balbianii,  118 

exneri,  121 

gaskrostei,  121 

heUandi,  121 

immersa,  119 

incurvata,  119 

sabrazesi,  120 
Sphaerospora,  10,  57,  100,  178,  185 
Sphaerospora  angulata,  102 

carassii,  103 

divergens,  100 

elegans,  100 

masovica,  101 

plaiessae,  102 

polymorpha,  102 

rostrata,  101 

sp.  Davis,  103 

sp.  Southwell  et  Prashad,  103 
Sphaerosporea,  10,  57,  86, 178 


503] 


STUDIES  ON  MYXOSPORIDIA—KUDO 


265 


Sphaerosporidae,  10,  57,  99,  178 
Spleen,  Myxosporidia  found  in,  41 
Spore,  description  of,  47 
Sporoplasm,  51 

Stomach,  Myxosporidia  parasitic  in,  39 
Sutural  diameter,  51 

edge,  51 

line,  51 

plane,  51 

ridge,  51 
Switzerland,  Myxosporidia  of,  23 
Tail  of  spore,  51 

Testis,  Myxosporidia  parasitic  in,  41 
Thickness  of  spore,  51 


Tissue,  Myxosporidia  parasitic  in,  37,  43,  45 

Trophozoite,  51 

United  States,  Myxosporidia  of,  15 

Urinary  bladder,  Myxosporidia  parasitic  in, 

41 
Vegetative  form,  51 
Wardia,  9,  56,  82 
Wardia  ohlmackeri,  83 

ovinocua,  82 
Zschokkella,  10,  58,  122 
Zschokkella  acheiiognalhi,  123 

globulosa,  123 

hildae,  122 

nova,  122 


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